Puffballs and their allies in Michigan.
Smith, Alexander Hanchett, 1904-

Page  i PUFFBALLS AND THEIR ALLIES IN MICHIGAN

Page  ii

Page  iii PUFFBALLS AND THEIR ALLIES IN MICHIGAN ALEXANDER H. SMITH ANN ARBOR University of Michigan Press I95

Page  iv Copyright I95I By the University of Michigan

Page  v CONTENTS PAGE INTRODUCTION........................................... 1 ACKNOWLEDGMENTS...................................... 5 THE FRUCTIFICATION..................................... 6 EDIBILITY............................................... 11 KEY TO ORDERS OF GASTEROMYCETES........................ 12 HYM ENOGASTRALES....................................... 13 Key to Families of Hymenogastrales.................... 13 G asterellaceae....................................... 13 Key to Genera of Gasterellaceae.................... 13 Gasterella Zeller and W alker...................... 14 Gasterellopsis Routien........................... 14 M elanogasteraceae................................... 15 Key to Genera of Melanogasteraceae................ 15 M elanogaster Corda............................. 15 A lpova D odge................................... 17 Rhizopogonaceae.................................... 18 Key to Genera of Rhizopogonaceae................. 18 Rhizopogon Fries................................ 18 Hymenogasteraceae.................................. 20 Key to Genera of Hymenogasteraceae............... 20 Hymenogaster Vittadini.......................... 20 H ydnangiaceae...................................... 21 Key to Genera of Hydnangiaceae................... 21 Arcangeliella Cavara............................. 22 GAUTIERIALES..................................... 23 G autieriaceae....................................... 23 Gautieria Vittadini.............................. 23 TREMELLOGASTRALES..................................... 25 Key to Families of Tremellogastrales.................... 25 Tremellogasteraceae................................. 25 Key to Genera of Tremellogasteraceae.............. 25 G astrosporiaceae.................................... 25 Gastrosporium Zeller............................. 25 H YSTERANGIALES......................................... 26 Key to Families of Hysterangiales.....................2. V

Page  vi vi PUFFBALLS AND ALLIES IN MICHIGAN PAGE Hysterangiaceae..................................... 26 Key to Genera of Hysterangiaceae.................. 26 Hysterangium Vittadini.......................... 27 Phallogaster M organ............................. 28 Protophallaceae..................................... 29 Key to Genera of Protophallaceae.................. 29 Gelopellaceae...................................... 29 Gelopellis Zeller................................. 29 PHALLALES.............................................. 29 Phallaceae.......................................... 30 Key to Michigan Genera of Phallaceae.............. 30 Mutinus Fries................................... 30 Key to Species of Mlit7innts......................... 31 Phallus Persoon................................. 33 Key to Michigan Species of Phallus................. 33 Dictyophora Desvaux............................ 34 LY^ COPERDALES......................................... 36 Key to Families of Lycoperdalel s........................ 36 Arachniaceae...................................... 37 Key to Genera of Arachniaceae..................... 37 Arachnion Schweinitz............................37 Lycoperdaceae...................................... 38 Key to Genera of Lycoperdaceae................... 38 Calvatia Fries...................................39 Key to Species of Calvatia......................... 39 Lycoperdon Persoon............................. 47 Key to Species of Lycoperdon7...................... 47 Disciscda Czerniaiev............................. 68 Key to Species of Disciseda........................ 69 Bovistella Morgan............................... 72 Key to Species of Bovistella........................ 72 Bovista Persoon................................. 75 Key to Species of Bovista.......................... 75 Broomeiaceae...................................... 79 Key to Genera of Broomeiaceae....................79 Mycenastraceae..................................... 79 Key to Genera of Mlycenastraceae..................79 Mycenastrnl I)esvaux........................... 80

Page  vii CONTENTS vii PAGE M esophelliaceae..................................... 81 Key to Genera of Mesophelliaceae................. 8 Geastraceae......................................... 81 Key to Genera of Geastraceae...................... 82 Myriostoma Desvaux............................. 82 Geastrum Persoon............................... 83 Key to Species of Geastrum....................... 83 SCLERODERMATALES....................................... 97 Key to Families of Sclerodermatales..................... 97 Sclerodermataceae................................... 98 Key to Genera of Sclerodermataceae................ 98 Scleroderma Persoon............................. 98 Key to Michigan Species of Scleroderma............. 99 Sedeculaceae........................................ 104 Sedecula Zeller.................................. 104 Pisolithaceae...................................... 105 Key to Genera of Pisolithaceae..................... 105 Pisolithus Albertini and Schweinitz................. 105 Glischrodermataceae................................. 1o6 Glischroderma Zeller............................. 106 Tulostomataceae................................... 1o6 Key to Genera of Tulostomataceae................ 107 Tulostoma Persoon.............................. 107 Key to Michigan Species of Tulostoma.......... 107 A straeaceae......................................... 113 Astraeus M organ................................ 113 Calostomataceae.....................................116 Calostom a Zeller................................ 116 NIDULARIALES........................................... 16 Key to Families of Nidulariales....................... 16 N idulariaceae....................................... 117 Key to Genera of Nidulariaceae..................117 Cyathus Persoon............................... 117 Key to Michigan Species of Cyathus................ 117 Crucibulum Tulasne............................. 120 Sphaerobolaceae..................................... 121 Key to Genera of Sphaerobolaceae.................121 Sphaerobolus Persoon........................... 121

Page  viii viii PUFFBALLS AND ALLIES IN MICHIGAN PAGE PODAXALES.............................................. 122 Key to Families of Podaxales........................... 122 Secotiaceae......................................... 122 Key to Genera of Secotiaceae...................... 123 Secotium Kunze................................. 123 Podaxaceae......................................... 125 Key to Genera of Podaxaceae...................... 125 LITERATURE CITED....................................... 125 INDEX.................................................. 129

Page  ix LIST OF PLATES (Plates I-XLIII follow page 131) PLATE I. FIG. 1. Melanogaster Broomeianus Berkeley in Tulasne. FIG. 2. Alpova cinnamomeus Dodge. FIG. 3. Gautieria morchelliformis Vittadini. II. FIG. 1. Phallogaster saccatus Morgan. FIG. 2. Mutinus caninus var. caninus. III. FIG. 1. Mutinus caninus var. albus Zeller. FIG. 2. Dictyophora duplicata (Bosc.) Fischer. IV. Phallus Ravenelii Berkeley and Curtis. V. Dictyophora duplicata (Bosc.) Fischer. VI. FIG. 1. Calvatia cyathiformis (Bosc.) Morgan. FIG. 2. Calvatia fragilis (Vitt.) Morgan. VII. Calvatia gigantea (Pers.) Lloyd. VIII. Calvatia craniformis (Schw.) Fries. IX. Calvatia saccata var. elata (Massee) Hollos. X. FIG. 1. Lycoperdon subincarnatum Peck. FIG. 2. Lycoperdon pusillum Persoon. FIG. 3. Lycoperdon oblongisporum Berkeley and Curtis. XI. FIG. i. Lycoperdon marginatum Vittadini. FIG. 2. Lycoperdon marginatum Vittadini. XII. FIG. 1. Lycoperdon pyriforme Persoon. FIG. 2. Lycoperdon pyriforme Persoon. XIII. Lycoperdon perlatum Persoon. XIV. Lycoperdon perlatum Persoon. XV. FIG. i. Lycoperdon perlatum Persoon. FIG. 2. Lycoperdon pedicellatum Peck. XVI. FIG. i. Lycoperdon umbrinum Persoon var. umbrinum. FIG. 2. Lycoperdon umbrinum var. atropurpureum (Vitt.) Holl6s. XVII. Lycoperdon umbrinum var. floccosum Lloyd. XVIII. Lycoperdon umbrinum var. floccosum Lloyd. XIX. FIG. 1. Lycoperdon rimulatum Peck. FIG. 2. Lycoperdon pulcherrimum Berkeley and Curtis ix

Page  x x PUFFBALLS AND ALLIES IN MICHIGAN PLATE XX. FIG. 1. Lycoperdon echinatuml Persoon. FIG. 2. Lycoperdon echinatum Persoon. XXI. FIG. 1. Disciseda candida (Schw.) Lloyd. FIG. 2. Disciseda subterranea (Pk.) Coker and Couch. XXII. Bovistella radicata (Mont.) Patouillard. XXIII. FIG. 1. Bovistella radicata (Mont.) Patouillard. FIG. 2. Bovistella radicata (Mont.) Patouillard. XXIV. FIG. 1. Bovista pila Berkeley and Curtis. FIG. 2. Bovista plumbea Persoon. FIG. 3. Bovista minor Morgan. XXV. FIG. 1. Mycenastrum corium (Guers.) Desvaux. FIG. 2. Mycenastrum COtrium (Guers.) Desvaux. XXVI. FIG. 1. Mycenastrum corium (Guers.) Desvaux. FIG. 2. Geastrum coronaturn Persoon. XXVII. FIG. 1. Geastrum coronaturn Persoon. FIG. 2. Geastrum pectinatum Persoon. FIGS. 3-4. Geastrum Schmidelli Vittadini. XXVIII. FIG. 1. Geastrum triplex Junghuhn. FIG. 2. Geastrum triplex Junghuhn. XXIX. FIG. i. Geastrum rufescens Persoon. FIG. 2. Geastrum rufescens Persoon. XXX. FIG. i. Geastrum saccatum Fries. FIG. 2. Geastrum saccatum Fries. XXXI. FIG. 1. Geastrum campestre (Morg.) Kambly and FIG. 2. Geastrum umbilicatum Fries sensu Morg; FIG. 3. Geastrum 1Morganii Lloyd. XXXII. FIG. 1. Scleroderma aurantilum Persoon. FIG. 2. Geastrum limbatum sensu Coker and Cot FIG. 3. Geastrum limbatum. XXXIII. Scleroderma aurantium Persoon. XXXIV. FIG. i. Scleroderma flavidum Ellis and Everhart. FIG. 2. Scleroderma flavidum Ellis and Everhart. XXXV. FIG. 1. Scleroderma arenicola Zeller. FIG. 2. Scleroderma lycoperdoides Schweinitz. XXXVI. Scleroderma Geaster Fries. XXXVII. Pisolithus tinctorius (Pers.) Coker and Couch. LXXVIII. FIG. 1. Tulostoma striatum Cunningham. FIG. 2. Tulostoma fibrillosum White. Lee. an. ich. X

Page  xi LIST OF PLATES xi PLATE XXXIX. FIG. 1. Tulostoma campestre Morgan. FIG. 2. Tulostoma simulans Lloyd. FIG. 3. Tulostoma simulans Lloyd. XL. FIG. 1. Astraeus hygrometricus (Pers.) Morgan. FIG. 2. Crucibulum levis (D.C.) Kambly. XLI. Crucibulum levis (D.C) Kambly. XLII. FIG. 1. Sphaerobolus stellatus Persoon. FIG. 2. Sphaerobolus stellatus Persoon. XLIII. Secotium agaricoides (Czern.) Holl6s.

Page  xii

Page  1 INTRODUCTION The group treated here, technically known as the Gasteromycetes, is one which always attracts the attention of nature lovers as well as the mycologist. Puffballs are encountered throughout the state in all seasons of the year. During the summer and autumn months the fresh immature fructifications are often used for food. During late autumn, winter, and early spring the old dried spore cases are objects of curiosity because of the manner in which they puff or send out a cloud of spores when kicked or stepped upon. For some reason, however, no one previously has made a serious attempt to organize the available information on the Michigan flora. This study was sponsored jointly by the Herbarium of the University of Michigan and the Cranbrook Institute of Science, Bloomfield Hills, Michigan. The aim was to produce an authoritative account of the Michigan puffball flora, which would be of use to the people of the state generally rather than to specialists alone. Since joining the staff of the University Herbarium in 1933, I have been collecting in this group, at first in a haphazard manner, but during the last five years intensively, whenever a favorable opportunity presented. Collecting these fungi, though not as difficult as collecting species of the Agaricaceae, is still not so easy that one can plan to do work at a certain time and actually accomplish it then. Puffballs are like mushrooms in that they fruit more or less at the whim of the weather, but unlike mushrooms their mature spore cases are mostly persistent and can be collected over a rather long period of time. Some will even overwinter. It might be worth mentioning in this connection that the study of mushrooms requires in most instances a different approach from the study of puffballs. The mushroom collector searches for the buttons and the expanding and freshly mature fruiting bodies; he is only secondarily interested in those past maturity. IL

Page  2 2 PUFFBALLS AND ALLIES IN MICHIGAN The student of the puffballs has little use for the immature fruiting body; he is concerned mainly with wTell-matured specimens, since those which are immature give him little information of value in identifying the species. One collecting fungi for food prefers young specimens regardless of whether they be mushrooms or puffballs. Tlhe reasons for this difference in approach are obvious, but are somewhat disturbing to one like myself, who took up tlle study of puffballs after having specialized in a study of the gill fungi. In botlh groups a proper evaluation of spore characters is essential to an understanding of speciation and the characterization of genera. In the n!mslhrooms spore deposits are obtained best from fruiting bodies jlst reaching maturity. In tle puffballs spore cliaracters are not usually reliable until the fruiting body is well matured or even past maturity. Since tlle spores of mushroomns are discilarged from the basidia and are produced over a slort period of time, such as a week to ten days in fleshy species, information about them can be obtained only wllile the carpoplhore is still fresh. The spores of puffballs are produced on basidia, but are not discharged. Tlie basidia eventually disintegrate, leaving the spores as a large mass inside tlhe spore case. The spores are eventually set free by tle opening of the spore case in whatever manner is clharacteristic for the particular species. The taxonomic claracters used to identify puffballs are cliefly tliose concerned with the wall of the spore case (called the peridium), its markings and the way it opens to release the spores, as well as with the characters of tlhe spores themselves and of any sterile hyphae or cells mixed in with them. All these are characters which can only be observed on mature fruiting bodies. Even observations of the markings of the outer wall of tlhe spore case must be ladce on mature specimens to give the cllaracters any value taxonomically. With thlis in mind it is easy to understand how much of tlle work on puffballs can be done in the laloratory from dried specimens on which few if any field notes lhave been taken. The mushroom specialist, however, often must havc notes on the characters taken from the

Page  3 INTRODUCTION 3 fresh fruiting body in order to identify the specimen. As every naturalist knows, each group of organisms develops according to its own pattern, and classifications and techniques to be good must be adjusted in each case to the pattern of the group in question. To return to the status of knowledge concerning the Gasteromycetes of Michigan, it should be pointed out that the present work is by no means a final report. It is offered in the hope that it will stimulate interest on the part of collectors in the state to explore more fully their respective communities so that we eventually may obtain information as to relative abundance of the commoner species, discover more stations for those which are rare, and add materially to the list of those known to occur here. Michigan is an ideal state for a floristic study such as this because of the diversity of its associations of higller plants and its varied topography. It is well known, for example, that certain of the higher plants common along the Pacific coast occur in the Upper Peninsula. It is probable that one may find a similar distribution also among the simpler Gasteromycetes. This opinion is based in part on the occurrence of Al[lova both on Isle Royale, Keweenaw County, and west of the Cascade Mountains. In fact the species of the order to which it belongs, the Hymenogastrales, or so-called basidiomycetous truffles, are still so poorly known in Michigan that students cannot lay claim to having made even a good preliminary survey of thein. During a warm wet season the forests of red and jack pine should yield a rich harvest in this group alone. In such large genera as Lycopcrdon and Geastrum, much work of a different sort remains to be done. Here extensive collections are needed to clarify species concepts and to obtain information on the variation caused by habitat, as is to be seen for instance, in species of Sclerodermna which grow both on sand dunes and on firmer substrata. In Lycoperdon I have deliberately been conservative in regard to the number of species recognized. When ample collections are available a number of interesting additions will certainly be made. The same may be said

Page  4 4 PUFFBALLS AND ALLIES IN MICHIGAN of Geastrum. Tulostoma presents a problem of exploration. The number of species recorded here can hardly represent the total Tulostoma flora of Michigan. No doubt a great deal of the lack of interest in bringing out a systematic account of the Michigan Gasteromycetes can be laid at the door of C. G. Lloyd, the well-known specialist from Ohio, who dominated this branch of mycology from about go1900 until after the first world war. Lloyd did not work on a regional basis. It seems clear to me that his great industry and the broad scope of his activities encouraged other mycologists to concentrate on other groups of fungi. In a survey of the literature his work, however, should be mentioned first. Although voluminous, it is not organized in such a manner as to make it convenient to use. In addition, it now appears (see Cunningham, 1942) that Lloyd himself was guilty of making many of the same mistakes that he so severely criticized in others. The publication by Coker and Couch on the Gasteromycetes of the eastern United States and Canada (1928) is indispensable to the serious student of this group. It is well illustrated and contains a wealth of information. But the authors did not use the code of nomenclature (International Rules of Botanical Nomenclature) which has since been accepted by most American workers and which I have followed here, so that differences in the names used for some species are to be expected. Cunningham's (1942) work on the species of Australia and New Zealand, though not generally available to people in this area, is one I strongly recommend to all who are interested in a general study of Gasteromycetes from the standpoint of relationships as well as floristics. It is conservative, however, and the student of the North American species will not find all of his questions answered there. Apparently, as in the case of the gill fungi, North America has one of the most diverse Gasteromycete floras of any region known so far. Consequently, students are looking forward to the forthcoming publication of the work on the North American species by our best-known specialist, the late Dr. S. M. Zeller. His classification was published in Mycologia (1949) in the form

Page  5 A CKNO WLEDGMENTS 5 of keys to the orders, families, and genera, together with descriptions of the orders and families. I have used his outline in this work. The quoted descriptions of orders and families are the ones published in Mycologia. His keys have been adopted with minor changes in arrangement of choices and wording. In view of the probability that many more Gasteromycetes will be found in Michigan than are included in this work, I think that the fairly complete keys to genera will prove very useful. Zeller's classification, it is true, needs to be tested. His recognition of the family Astraeaceae as a member of the Sclerodermatales will doubtless bring forth some strong criticism. To decide this problem properly a detailed study of Astraeus Pteridis is needed in addition to what is already known of A. hygrometricus. Johnson (1929) published an account of the Gasteromycetes of Ohio, and Kambly and Lee (1936) made a similar study for Iowa. Both these publications contain accounts of species which have not yet been reported from Michigan but which will probably be found here. In Michigan, Longyear (1904), then at Michigan State College at East Lansing, published the first reasonably extensive list of species, and later Kauffman (1908) published a list based on his own collection. Swartz (1933), one of Kauffman's students, did considerable collecting in the vicinity of Ann Arbor in connection with his studies on the development of certain species. Routien (1939, 1940) collected and described a number of species from near East Lansing, which are both very minute and very interesting. ACKNOWLEDGMENTS It is a pleasure to acknowledge the assistance and helpful criticism of those who have contributed to the completion of this work. Dr. E. B. Mains, Director of the University Herbarium, has contributed collections and photographs in addition to critically reading the manuscript. Through the courtesy of Dr. Ernst E. Bessey, of Michigan State College, the collections in that institution were studied and some very interesting information obtained. The late Dr. S. M. Zeller contributed in

Page  6 6 PUFFBALLS AND ALLIES IN MICHIGAN formation on nomenclatorial problems and also identifications in several instances. I am indebted to Morten Lange for checking the microscopic characters of the Michigan collections in the University Herbarium. To Victor Potter, of Ithaca, Michigan, I am especially indebted for excellent collections from that area. They contain several new records for the state as well as abundant material of many rare species. THE FRUCTIFICATION Typically, tle fructification of the puffball plant is a reInarkably simple structure more or less globular in sllape. The name Gasteromycetes, whlich is applied to the group, means stomacl fungi. The mature fruiting body consists of a spore case filled with spores and often some sterile modified threads (hyphae) or parts of them. The mass of spores with the included sterile elements is known as the gleba and the surrounding wall or walls as the peridium (plural peridia). For an example of this type of fruiting body see the illustration of Bovista p)ila (PI. XXIV, Fig. i). The modifications wlicll this simple type of structure has undergone in the course of evolution are exceedingly diverse, and tile almost numberless adaptations exlibited by the members of the group make a fascinating study. One of the first deviations from thle simple type is the development of a stemlike base, in its final expression a true stipe, as in Tulostoma fibrillosulm (P1. XXXVIII, Fig. 2). This adaptation is found in all tle major groups, though in the Hymenogastrales (the basidiomycetous tubers and their relatives) it is rare. In the Phallales, the stalk is the structure (called tlie rcceptaculumll) whicll elevates the spore mass for dispersal, eitller on its apex or over tlhe surface of a distinct, apically attached pileus. In tle Sclerodermatales a structure known as tlhe pseudostipe (P1. XXXIII) develops. Tllis consists of a mass of fibrous myceliumn whiclh binds togethler tiglhtly so mucl soil and debris tlat when collected the specimens appear to have a stipe. This structure apparently serves chiefly for anchorage, since it is likely to reacll its best development on loose substrata such as sandy soil.

Page  7 THE FRUCTIFICATION 7 In the Lycoperdaceae, by definition, species with well-developed stipes are not included, but nevertheless within that family numerous stemlike adaptations occur, as the sterile base in Bovistella radicata (Pls. XXII, XXIII). A well-developed rootlike organ anchors the fructification. The sterile lower part of the fructification has a chambered structure that is known as the sterile base or subgleba. The subgleba apparently serves to elevate the spore mass (gleba) to a position more favorable to spore discharge. This type of a sterile base is a feature of many species of Calvatia and Lycoperdon, and tile degree of its development has been used to some extent in the delimitation of species. The family Tulostomaceae is distinguished from the Lycoperdaceae by the presence of a true stipe. The stipe may be mostly embedded in the substratum or seated on top of it. Members of this family are most numerous in arid regions such as the southwestern United States. In the Nidulariales a stipe is not present, though some of the fructifications may have a stipelike base. Evolutionary changes in this group have taken place chiefly in other lcharacters. Most of the adaptations developed by the Gasteromycetes have a bearing on the problem of spore dispersal, and the manner in which the fruiting body is constructed so as to facilitate spore dispersal, either by wind or by insects and animals, has always been given great emphasis in systems of classification. In the Hymenogastrales, here regarded as the most simplified members of the subclass Gasteromycetes, one finds no specialized means of spore discharge or liberation. Thl e fructification develops in the ground or humus just under the surface, but frequently becomes exposed by the time it is mature and then either decays or is eaten by rodents. Many species develop strong odors as the fructifications age, and these may serve to attract insects or rodents which will carry away spores on some part of their anatomy or in their digestive tracts. At best this seems to be a very poor method of dispersal, and one wonders low long it has taken certain of the species to attain tlhe circumpolar distribution which they are known to have.

Page  8 8 PUFFBALLS AND ALLIES IN MICHIGAN The highest degree of specialization for spore dispersal by insects has been reached in the Phallales. Here morphological specialization has accompanied specialization in the production of odors peculiarly attractive to insects, as in Dictyophora duplicata (P1. III, Fig. 2). IMutinus and Phallus represent different stages in the development of types. The former is simple because it lacks a pileus, and the latter is more complex because it possesses one. Phallogaster saccatus (P1. II, Fig. i), a memb)er of the Hysterangiales, has developed definite characters similar to those of the Phallales. Species of Phallales are most abundant in the tropical and warmer regions generally, and anyone interested in the evolution of types of fructification within the order should consult such works as those of Cunningham, Coker and Couch, and Lloyd. In I\ichigian representatives of the Pliallales the gleba has a cllaractcristically foul smell somewhat similar to that of decaying flesh. In addition, it is elevated on the elongated receptaculum to such a height that the odor becomes well dispersed and attracts insects from a relatively wide area. The slimy nature of the gleba ensures that spores will adhere to anything which touches it. In the Sclerodermatalcs are examples of relatively simple types of spore dispersal. In Scleroderma aurantium the wall of the spore case eventually fragments or at least opens by an irregullar apical rupture. In both S. flavidum and S. Geaster the wall segmnents in a fairly distinct pattern as is shown in the photographs (Pls. XXXIV and XXXVI). In Pisolithus the spores are produced in capsule-like structures which fragment, releasing the spores. All stages of the process are shown in Plate XXX\ II. In the Calostomlotaceae the spore case opens by an apical pore, and the fructification in other respects is more liiglily specialized. There are no representatives of it in Michigan, lbut at least one species is common southward. Some authors do not admnit this family to the Sclerodermatales. 'The Iycop:l'ralcs exhibit considerable diversity in the manner in which tlhe spores are shed. In Calvatia the method is very simple. At maturity the wall of the fructification breaks

Page  9 THE FRUCTIFICA TION 9 up into fragments, and as these fall away the gleba is exposed to the action of the weather. This type is illustrated by Calvatia giganlea (P1. VII). According to some authors the genus Calvatia is characterized by having a single peridial layer. Tle situation here is interesting from a scientific standpoint, but I do not emphasize it because in most species of Calvatia there appear to me to be two layers to the wall, an outer layer which usually cracks up into some sort of distinctive pattern and an inner membranous one. Lycopcrdonl is characterized by having a wall of two layers: the inner wall, or endoperidium, and the outer, or exoperidium, the latter being present as spines of varying size. The spores are shed through an apical pore. This genus is thus more highly specialized than Calvatia. In Mycenastrum, Bovista, and Bovistella the method of spore liberation is either by way of an apical irregular rupturing or by a segmenting of the peridium. One Michigan species of Disciseda opens by a pore at the base of the fructification instead of at the apex, as in Lycoperdon. When the outer wall splits in a belt around the middle, half of it remains as a cap over the apex of the spore case. The lower half adheres to the soil so that the endoperidium is exposed over the basal half and the pore forms at the original place of attachment. As the fructification is blown around the spores are shaken out. The most highly developed system for spore liberation in the Lycoperdales is in Geastrum. Here the exoperidium constists of three layers and opens by splitting into segments. The segments, called rays, bend backward and finally lift the spore case up into a more favorable position for spore dispersal. The spore sac, consisting of the endoperidium, exhibits considerable specialization in the type of apical pore formed. The formation may be indefinite, that is, the pore zone (also known as the peristome) may not be clearly marked off from the remainder of the endoperidium, or it may be definite, that is, marked off by a circular depression or ridge. Furthermore, the mouth, or pore, may be formed by closely arranged fibrillose segments each of which tapers to a point, or the segments may be grooved and be described as sulcate. All of these adaptations indicate a higher degree of specialization than exists

Page  10 10 PUFFBALLS AND ALLIES IN MICHIGAN in other members of the family Geastraceae. In Mesophellia, a genus not yet known to occur in Michigan, there is a threelayered exoperidium and an endoperidium somewhat as in Geastrum, but the fructifications never open up. As already pointed out, the family Tulostomaceae differs from the Lycoperdaceae by the presence of a stipe. This family has various degrees of specialization in the manner of liberating spores. In Schizostonma the spore case opens by irregular fissuring. In Tulostonma it opens by a pore which may be more or less distinctive for certain species. In Battarraea, a genus usually encountered in the warmer arid regions of the world, the apical half of the spore case separates by circumscissile cleavage and falls off leaving the gleba exposed. The Nidulariales present a totally different approach to the problem of scattering the spores. In this group the spores are forned within small bodies known as peridioles, which in turn are produced in a characteristic type of fructification. The whole often resembles a small nest with eggs in it, hence the common name "bird's-nest fungi." In some species these peridioles are forcibly discharged from the "nest"; in others they simply fall out eventually and unless eaten by some animal, never come to rest far from the original nest. The peridioles may germinate to give rise to new mycelia, or, depending upon the conditions, they may break down and eventually liberate the spores. As may be seen from this discussion of the stipe and of the methods of spore discharge, there are no clear-cut major lines of development. These characters have originated independently in many different groups and show varying amounts of progress. In some sections of the present classification the differences in these characters are given great emphasis; and in other places they are not, depending somewhat on the opinion of the investigator, but to a greater extent on the correlation of more fundamental characters, such as the spore type, the presence of capillitium, and the type of development of the fructification. Because my experience with the group is only for a restricted area, I do not feel qualified to discuss the evolution of the Gasteromycetes

Page  11 EDIBILITY 11 and the relationships of the various genera to each other. I have merely arranged the Michigan species according to the classification of Zeller (1949). EDIBILITY I have always discouraged, and doubtless will continue to discourage, people from eating the eggs of stinkhorns and the various species of Scleroderma. It must be admitted, however, that there are reports in the literature to indicate that these are edible. Mcllvaine and Macadam (1912) recorded that the eggs of a number of the phalloids which are included in this work may be eaten and that those of Phallus impudicus are "tender and agreeable food." This would appear to leave the Gasteromycetes as a group without any species known to cause serious poisoning regularly and certainly indicates that it is one of the safest groups of fungi for anyone to use. One should not, however, rely so heavily on their good reputation as to become careless. I have known people to have rather violent gastrointestinal upsets from eating species of Calvatia and Lycoperdon, two of the safest genera. Some are known generally to be rather violent purgatives. In some instances the trouble may be caused by eating specimens which are on the verge of maturity. One should be careful to use only those specimens which are white clear through. If discolorations have begun to develop it is likely that the flavor will be bad. Many who have tried puffballs and found them unpalatable may have eaten material that was old. The larger species of Calvatia, in particular Calvatia gigantea and, to a lesser extent, C. cyathiformis, are popular in southern Michigan and are frequently sold on farmers' markets in the fall. C. craliformis, when it can be found in quantity, is also deserving of mention. But anyone who has not eaten one of these species previously, even the two just mentioned, should try only a small portion the first time as a precaution. A person who eats C. gigantea regularly may have difficulty with one of the others, particularly if he eats very much the first time he tries it. It is to enable people to draw their own conclusions about the relative merits of the various species that I urge that

Page  12 12 PUFFBALLS AND ALLIES IN MICHIGAN they learn to recognize the species and try them one at a time. In this way one can build up a reliable fund of knowledge in regard to his own likes or idiosyncrasies. The members of the Hymenogastrales are edible as far as is known, and it is reported in the literature that a few species are eaten as a substitute for the true truffle. No doubt the lack of information on these fungi in North America has been largely due to the difficulty of finding and identifying them. The true truffles belong to the genus Tuber, and are ascomycetes, not basidiomycetes. They are even rarer in Michigan than are the members of the Hlymenogastrales. KEY TO ORDERS OF THE GASTEROMYCETES i. Gleba or spore mass held in place until near maturity, on the underside of a centrally stipitate cap (peridium); basidia at first in a true hymcnium........................................ PODAXALEs, page 122 i. N ot as in above choice............................................... 2 2. Original structure of glcba maintained until maturity; rarely stalked (mostly hypogeous species); peridium not dehiscing in a characteristic manner.......................................................... 3 2. Gleba becoming disorganized by maturity; peridium usually opening in a characteristic manner................................. 5 3. Gleba typically fleshy rather than cartilaginous; basidia in a true hymenium (except in Melanogastraceae)......... HYMENOASTRALES, page 13 3. Gleba and/or peridium cartilaginous or gelatinous or both............. 4 4. Spores with longitudinal ribs.......................GAUTIERIALES, page 23 4. Spores echinulate or sculptured.................REMELLOGASTRALES, page 25 4. Spores bacillar, ellipsoid, smooth; basidia phalloid..IIYSTERANGIALES, page 26 5. Gleba slimy and odoriferous, finally exposed by the extension of a spongy stemlike structure known as the receptacle........... PHALLALES, page 29 5. Glebal cavities isolated from each other (as hard or brittle egglike bodies or discs) by dissolution of tissue lying between them (the fructification reminiscent of a nest with eggs in it).............. NIDULARIALES, page 116 5. Gleba a powdery or pulpy mass at maturity (sandy to rough in Arachniaceae).......................................................... 6 6. Gleba with s)ymmeLrically distributed basidia or with basidia-bearing nests or lacunae arising through the dissolution of fundamental tissue, typically without a well-organized hymenium..SCLERODERMATALES, page 97 6. Gleba chambered by the outgrowth of tramal plates or pegs, walls of chambers covered with at least a rudimentary hymenium of basidia... 7 7. Peridium with 1 or more gelatinous layers......REMELLOGASTRALES, page 25 7. Peridium without gelatinous layers................. LYCOPERDALES, page 36

Page  13 HYMENOGASTRALES 13 HYMENOGASTRALES "Fructifications mostly hypogeous, bulblike, occasionally pear- or spindle-shaped, stalked, or epigeous; rarely with a stemlike columella; peridium remaining indehiscent to maturity, seldom disintegrating early; gleba of one or more cavities, lacunae filled with gel or with basidia-bearing hyphae or nests, holding original structure to maturity; peridium and gleba essentially fleshy, not cartilaginous; conidiophores when present borne in hymenium with basidia (Holocotylon) or in a separate fructification (Leucophleps, conidial stage of Leucogaster)" (Zeller, 1949: 38). KEY TO FAMILIES OF HYMENOGASTRALES i. Fructification minute, with a single glcbal cavity at maturity........... 2 i. N ot as above........................................................ 3 2. Spores smooth.......................................... PROT;GAs-rT RACEAE 2. Spores verrucose........................................ GASTERELLACEAE 3. Gleba with open cavities lined with a true hymenium.................. 4 3. Not as above, cavities filled with gel or lined with a false or rudimentary hymenium......................................... MELANOGASTERACEAE 4. Spores smooth..........................................RIIIZOPOGONACEAE 4. Spores verrucose.....................................HYMENOGASTERACEAE 4. Spores echinulate.........................................HYDNANGIACEAE GASTERELLACEAE "Fructifications very small, depressed globose, epigeous, campanulate development; gleba finally uniloculate, but at times with one circle of cavities formed by vertical, centripetal invaginations which reach the center forming a false columella; cavities lined with a basidial hymenium; spores verrucose, dark" (Zeller, 1949: 39). KEY TO GENERA OF GASTERELLACEAE 1. Gleba at first and at maturity consisting of a single, hymcnium-lined cavity....................................................... G asterella 2. Gleba at first with a percurrent columella around which is 1 circle of cavities formed by vertical centripetal invaginations which reach the center, forming a false columella, and break away again at maturity; finally uniloculate........................................ Gasterellopsis

Page  14 14 PUFFBALLS AND ALLIES IN AMICHIGAN Gastcrclla Zeller andl Walker Fructification minute, 200-700 p in diameter, subglobose; wall of simple fundamental tissue; indehiscent; sterile base pulvinate to broadly conic, pseudoparenchymatous; gleba unilocular; hymenium smooth; spores brown, citriform, apiculate, verrucose. Gasterella hutoplhila Zeller and Walker Fructification depressed globose, 200-700 p in diameter, white at first, becoming liglit brownish drab, surface dry, cottony to innately fibrillose; sterile base pulvinate to subconic, pseudoparenchymiatous; peridium of a simple layer of loosely interwoven hyphlae, mostly parallel with the surface, hyaline, about 20 Ip thick; gleba uniloculate, brown, cavity empty; hymenium smooth; basidia clavate, 2- to 4-spored, soon evanescent; cystidia capitate-clavate, tle head dark brown, verrucose like tle spores; spores broadly citriform, with a broad apiculus; exospore dark brown, uniformly verrtucose except the lighter-colored apiculus, 10-12X 12-14 p. Hlabit, haitat, (and distri)btioni.-Routienr ( 1!)9) reported tlle fungus from Mlichigan on soil brolught into the laboratory from a wooded area near East Lansing, Inghlam County. Discussioni. —The description of genus and species is adapted fro(m Zeller and 'Walker (1935). It is a species one is not likely to reco'gnize in the field. Gastcrcllopsis Routien Fructification small, subglobose; peridiumn of inflated cells, dehiscent in a circumscissle manner at base; gleba uniloculate, witlh vertical centripetal infoldings tlat usually reach to the coltumella; collumella central, percurrent; spores citriform, apiciulate, verrluc(os, brownish black. GastcrcIlop)Ssis.silicola Routien Fructification sphlerical to depressed, 1-2 mm. in diameter; white, then black; pcridiuml dehiscent at the base of the columella, dissolving at mnatlurity; tramal plates finally dissolving;

Page  15 HYMENO GAs TRALES 15 basi(lia clavate, 2- to 4-sporcd; sporcs 14.5 (6.5)- 8 x 11 (13)14.5 p, pedicels measuring 2 X 1.5 p. Habit, habitat, and distribution.-On soil brought into laboratory from a woods near East Lansing, Ingham County. Known only from a single locality. Discussion.-The descriptive data have all been adapted from Routien (1940). The fungus occupies a very interesting place in the series of developmental types occurring in the Gasteromycetes. MIELANOGASTERACEAE "Fructifications subglose, usually hypogeous, sometimes stipitate at maturity; gleba of lacunae with basidia in nests or in a rudimentary hymenium from the walls of jelly-filled or pseudoparenchyma-stuffed cavities; gleba not becoming powdery; capillitium none" (Zeller, 1949!: 39) KEY TO (;C:NFRA OF MI.ANO(ASTLRACEAE i. Gle)ba b)lack or brown, with basidia scattered through gel-filled lacilnac.. 2 i. Glcba light-colored to white............................... 4 2. Gleba black, marbled with wlitish veins..................... l1clatilo(nstr 2. Gleba dark brown, marlled with lightt-trown vcins.................... 3 3. Spores ellipsoid, sm ooth............................................. 4 / ova 3. Spores spherical, echinulate; lactiferous d(lcts in peilidiin and septa................................................................. 1(( rC(1 o n ia 3. Spores citriform, rough with ](ose cxos,;lore..................C/lol(drostcr 4. Fructification stipitate....................................... Torrcdia 4. N ot as above........................................................ 5 5. Gleba lacunate, not chambered, hard..........................Cord itu cra 5. Gleba chambered, irregular hylmeniul liningl clallll m rs filled with hyphal tissue or gel....................................................... 6 6. Spores borne on conidiophores............................... cucol l )s 6. Spores borne on basidia.............................................. 7 7. Spores ellipsoid, smooth.............................. c...... cogastcr 7. Spores mostly spherical, ornamented or with a gelatinous sheath.................................................................. Iluc(a'tslc'r Mlelanlogastcr Corda Fructification hypogeous but by maturity often breaking through the soil, globose to irregular in shape; surface covered with branched rhizomorphs arising from exterior of wall and usually most numerous at base; peridilum a single layer of woven\

Page  16 16 PUFFBALLS AND ALLIES IN MICHIGAN gelatinized hlyphae, in the interior continuous with tranial plates; gleba black but marbled with whitish veins, of anastomosed traimal plates forming vesiculose to angular cavities which are filled with spores at maturity; columella absent; basidia irregularly distributed through a broad hyphal zone lining the cavities; spores dark-colored, smooth. Melaiogastcr amb iguus (Vitt.) Tulasne "Fructifications subspherical to ellipsoidal, 1-3.5 cm. in diameter, pale olive to olivaceous-brown when fresh, becoming cinnamon-brown to Prout's brown on drying, surface cottonyfurfuraceous, fibrils scanty, inconspicuous; peridium 300-6o00 thick when fresh, somewhat thinner in old dry material, hyaline within, yellowish-brown toward surface, prosenchymatous, lhomogeneous, with outer filaments fraying out to form the more or less erect, brown tomcntellum at the surface, of large, thickwalled, agglutinated hyphae which often collapse on drying; gleba black with yellowish septa, fundamental tramal tissue of more slender, smaller-celled prosenchyma, cavities filled with spores embedded in a gel; basidia clavate, evanescent, mostly 4-spored; spores (8-) 12-i6(-17) x (5.5-)6-8 (-8.5), obovoid with more or less acute apex, approaching citriform, thickwalled, smooth, very often uniguttulate, dark brown when mature" (Zeller and Dodge, 1937b: 642). Habit, habitat, and distribution.-Under hardwoods from April to October. The single Mlichigan collection was made by A. H. W. Povah, on July 29, 1927, near Ann Arbor, Washtenaw Colunty. Mclan ogastcr Broomcianus Berkeley in Tulasne (P1. I, Fig. i) "Fructifications subglobose to irregularly lobed, often coalescing when cespitose, 2-4 cm. in diameter; surface ochraceous then Dresden brown with a flesh tint, becoming blackish where handled, drying tawny-olive to warm sepia, appressedly tomentulose; brownish fibrils below; peridium variable in thickness, 50-200 p thick (mostly 5o-loo p), of stupose prosenchyma with

Page  17 HYMENOGASTRALES 17 large vesiculose cells, yellow-amber to darker toward the exterior, extending as fundamental tissue into the trama; gleba sooty-black, gelified, tramal septiments white to creamy, of subgelatinous hyaline hyphae, cavities filled with spores in a gel; spores light brown, narrowly oblong, broadly truncate below, (5-)6-8.4(-1 1) x (2.5-)3.5-4.0 (-4.5) P; odor of coffee grounds" (Zeller and Dodge, 1937b: 647). Habit, habitat, and distribution.-Single to gregarious under hardwoods. The collections of Kauffman, Ekwall, and Smith, as cited by Zeller and Dodge, are the only specimens I have seen. It fruits in Michigan during late summer and fall. Discussion.-According to my experience this is one of the species of Melanogaster most frequently found in the state, but it is nevertheless rare. A lpova Dodge Fructification oblong to subglobose; peridium single; gleba gelatinous, cavities soon filled; basidia irregularly arranged but evenly distributed in the cavities; spores hyaline under microscope, brownish in mass, ellipsoid, smooth; columella absent; mycelium attached to fructification in many places but inconspicuous and at maturity usually gelatinized. Alpova cil lamnozicus Dodge (P1. I, Fig. 2) Fructification oblong to globose, 4-20 mm. in diameter, whitish then pinkish buff to cinnamon buff, turning hazel to auburn (Ridgway); peridium thick (about 300 p), composed of large-celled pseudoparenchyma; gleba clay color, turning Hessian brown, gelatinous, the spaces between the septa at first filled with large spherical cells which finally disintegrate; septa of large, thin-walled, hyaline, parallel hypliac which simulate pseudoparenchyma; basidia on slender funiculi as in the Podaxaceae, scattered irregularly in the gel, very long and slender, 20-22 X4-5 I, 8-spored witl sterigmata about i \ long; spores hyaline under the microscope, pale brown in mass, ellipsoidal, 3-4 x 1.5-2.5 P.

Page  18 18 PUFFBALLS AND ALLIES IN MICHIGAN Habit, habitat, and distribution.-Half buried in the soil, often under alders (Alins), June 30 to September. Known in Michigan only from Isle Royale, Keweenaw County. Discussion.-Tlhis interesting genus was first discovered on Mount Hood, Oregon, by Wehmeyer and Kauffman, but it was described by Dodge (1931) from material collected on Isle Royale l)y C. A. Brown and A. H. IV. Povah. The species was named in honor of Dr. Povah, one of the members of the University of Michigan's expedition to the island in )930o. Since its discovery tile funguls las been reported as much more abundant in Washington, Oregon, and California than in Miclliian, and Zeller (1939) lias made a detailed study of its development. He found it mostly under conifers. My own collections along the West Coast confirm his observations. RHIZOPOGONACLAL "Fructifications subglobose, lypogeous or epigeous; peridium simple, witl or without rlizomorplic fibrils over the surface; gleba fleshy, not cartilaginous, witli open irregutlarly arranged cavities, or cavities diverging from the base or from a branched or simple colnumella; original structure of gleba maintained to maturity; basidia in true h1ymienia; spores smooth, tinted" (Zeller, 1949: 40). KEY TO (;ENYIRA OF RIIIZOPO(;ONACLAE i. Fructification typically hypogcous, with rhizomorphic fibrils over surface of lperidium; hymeniulli of basidia and paraphyses.............. 2 i. Not as abo\e........................................................ 3 2. (;]Cl)a willout colllulIlla ()r cons])icuo)us dendroid tran- al plaltes.. llhizol ogolz 2. Gleba with colunlllla or consll)iCUOUS dendroid tramal plates. Truncocolumlclla 3. Fructification epigcous, witli no superficial fibrils, hymenlium of basidia, conidiophores, and paraph ycs...............................Ho yloon 3. Fructification wlitli 11ollo0 stem; elieous; spores ellipsoid, smooth.. Lc Ratia Rhlizopo(goi Fries Fructification var-yilng from globose to tuber-like; peridium toutill, consistilng of( st ttpose and sometimes gelatinized hyplae arranged in 1 to 2 layers: surface covered with few to many laterally adliering rliizomorphs; gleba of persistent tramal plates

Page  19 HYMENOGASTRALES 19 anastomosed to form subglobose to labyrinthiform cavities; columella absent; spores hyaline to slightly tinted, smooth. Rhizopogon rubescens Tulasne "Fructifications cespitose, ovate, or irregularly globose, 1-6 cm. in diameter when fresh, 1-5 cm. when dry, color white at first, then livid yellow, reddening in air (Tulasne), and drying Morocco red to claret-brown or almost black where touched; odor weak or almost none (Tulasne); fibrils inconspicuous, innate-appressed above, simple, large, rlizomorph-like below, at first white (Tulasne), then reddening and becoming almost black; peridium thin, about 160-220 1, simplex, compact, brittle, very dark tawny; gleba at first white (Tulasne), then melleus to Isabella-color, brittle; cavities subglobose to labyrinthiform and irregularly crowded, empty; septa narrow, about 40-50 p broad, hyaline, usually not scissile until old; basidia pyriform or clavate, 2-8-spored, 12-14 P long; sterigmata about as long as ttle spores; spores acrogenous, oclhraceous-tawny in mass, lyaline or cream-colored under the microscope, ellipsoidal, 5-10 X 3-4.5 P, 1-2-guttulate, smooth" (Zeller and Dodge, 19i18: 1(). Habit, habitat, and distributtion.-In sand under pines. D. V. Baxter collected it near Ann Arbor, Washtenaw County, in a nursery under pine on October 17, 1927. The collection was identified by Zeller. Discussion.-The fructifications of the Michigan collection are all small. Rhizopogonl roscolus (Corda) Zeller and Dodge "Fructifications globose to irregular, 1.5-3 cm. in diameter, cinnamon-buff to Verona brown and even blackening on drying; fibrils scanty or disappearing, innate-appressed, black when dry; peridium thin, 160-300 p thick, compact, tawny under tile microscope; gleba from warm buff to bucktlorn-brown when dry, brittle; cavities subglobose and folded to labyrinthiform, empty; septa about 100 p broad, made up of closely woven, branching, hyaline hyphae witl tlhick gelatinized walls, not scissile; basidia ellipsoid, 12-13x7 [1, with small-lumened, heav

Page  20 20 PUFFBALLS AND ALLIES IN MICHIGAN ily gelatinized walls, mostly I-2-spored, seldom 3-5; sterigmata 10-14 p long, spores oblong to ellipsoid, acrogenous, dilute cream-colored under the microscope, heavy-walled, smooth, 2 -guttulate, with an equatorially placed nucleus, making the spores appear 1-septate, 8-12X3-5.5 p" (Zeller and Dodge, 1918: 16). Habit, habitat, and distribution.-Kauffman made 1 collection near Saginaw Forest, Ann Arbor, Washtenaw County, October 2, 1926. It was identified by Zeller. I have 2 collections made in October under pine at Proud Lake, Oakland County. Discussion.-Most species of this family have not been collected sufficiently to establish their distribution in the state. HYMENOGASTERACEAE "Fructifications globose, hypogeous, with or without rhizomorphic fibrils over the surface of the peridium; peridium simple; gleba fleshy, not cartilaginous, dark, with or without columella; with true basidial hymenium; spores dark, verrucose" (Zeller, 1949: 40). KEY TO GENERA OF IIYMENOGASTERACEAE i. Fructification with dendroid of percurrent columella; spores verrucose, mostly citriform.........................................Gymnoglossurn 1. Fructification without colunella...................................... 2 2. Spores verrucose, mostly citriform...........................Hymenogaster 2. Spores angular (as in Rhodophyllus of the Agaricaceac)..........Richoniella Hymenogaster Vittadini Fructification globular, pear-shaped to tuber-like, attached by a short radicating base or by rhizomorphs; peridium 1- to 2-layered, of stupose hyphae or pseudoparenchyma; gleba of tramal plates anastomosed to enclose subglobose cavities; hymenium lining the cavities as a palisade; columella absent; spores colored, verrucose or with a wrinkled gelatinous exospore. Hymenogaster niveus Vittadini "Fructifications about 1 cm. in diameter, subspherical, drying somewhat irregular, snow-white, reddening to the touch

Page  21 HYMENOGASTRALES 21 when fresh, drying ochraceous-tawny to buckthorn-brown or even darker, sterile base evident; peridium 300-320 p thick, drying 6o-1 lo p, composed of compactly woven hyphae (prosenchyma in islands) with larger varicose hyphae 5-6 p in diameter; gleba light fuliginous, becoming very dark brown, cavities large; septa thin, 15-20 p thick, composed of slender, compact, gelified hyplae; basidia 2-spored, pyriform, 8-9 x 5-6 p, projecting above the septa, not numerous and early collapsing, so that the sterigmata appear to arise directly from the septal hyphae; spores rather large, warted, short-pedicellate, ovoid- to ellipsoid-citriform, with short, blunt, hyaline apiculus, 15-18.5 x 9.5-11.5 p. Odor of Pelargoniinum" (Dodge and Zeller, 1934: 654-55). Habit, habitat, and distribution.-Growing under the surface of the soil (hypogeous) in forests of deciduous and coniferous trees. Known in Michigan from 1 collection, made by C. H. Kauffman, October 31, 1926, in Saginaw Forest, near Ann Arbor, AWashtenaw County. Discussion.-Since Dodge and Zeller recognized 54 species in their monograph, a careful searchl in Michigan should lead to the discovery of more than the single one included here. HYDNANGIACEAE "Fructifications hypogcous, subglobose or subpileate; with or without a columella or stipe, with campanulate development; gleba fleshy, not cartilaginous, maintaining original structure to maturity; with true basidial hymenium; sometimes with lactiferous ducts; spores echinulate, slightly tinted, usually spherical" (Zeller, 1949: 40-41). KEY TO GENERA OF IIYDNANGIACEAE i. Gleba with columella; lactiferous ducts in some or all tissues...Arcangcliclla 1. N ot as above...................................................... 2 2. Gleba with columella or stipe; tissues lacking lactiferous ducts but always with islands of pscudoplarnclhynla.........................Elasiomyces 2. Columella lacking, or tissues without lactiferous ducts; spores subglobose. 3 3. Spores thick-walled; glela gelatinous, cavities filled with spores...Sclcrogasler 3. Spores with thinner walls; gleba fleshy, not gelatinous..........Hydnanlgiu

Page  22 22 PUFFBALLS AND ALLIES IN MICHIGAN Arcangcliclla Cavara Fructification gregarious, hypogeous or emergent, fleshy, lactiferous; peridium thin, especially below, where it reaches the base in young specimens, often evanescent at maturity; columella usually percurrent, sometimes branched; base more or less sterile, ustually becoming attenuated into a stipelike attachmnent to the rhizonlorphs, generally lactiferous; gleba fragile or cheesy, lactiferous, cavities variable in size, often minute, irregular, radiating more or less from the columella and base; basidia 2- to 4-spored; cystidia sometimes present; spores spherical to ellipsoidal, eclhinulate to verrucose, often alveolate or somewhat eticulate. Arcangcliclla astcrosj)crma (Vitt.) Zeller and Dodge "Fructifications subspherical to reniform, up to 5 x3.5 cm.; surface tubercular-verrucose, pulverulent, with numerous fibrils, pure white with pinkish flecks which disappear, becoming greenish passing to deep olive, dirty brown to the touch, avellaneous or warm sepia to wood-brown or bister in alcohol, drying cinnamon-buff to sepia or even black; sterile base pulvinate to palmate in vertical section, prosenchymatous with hyaline lactifcrous ducts, on drying prosenchyma partially collapsing; peridium 320-480 p thick, drying 150-200 p, hyphae with vesiculose cells on the outside, within prosenchyma of large, vesiculose cells and relatively few lactiferous ducts; gleba white, exuding a salmon-tinted milk. brownish turning blue-black on exposure, becoming warm sepia in alcohol, and drying cinnamon-brown to Prout's brown or snuff-brown; cavities comparatively large, more or less radially arranged, filled with spores; septa hyaline, prosenchymatous, of large, vesiculose cells often appearing pseudoparenchymatous, with lactiferous ducts, the larger, platelike septa 110-150 p (drying 75-100 p) thick, the smaller septa 3(-35 p\ (drying 20-25 p) tlick, basidia short, clavate, 20-22 p long, 4-spored, sterigmata 5-6 p long; cystidia fusiform, 50-60 X 13-14,J; spores spherical, thickly covered with large conical to pyramidal spines 2 p lolng, 13-16 p in diameter including the

Page  23 GA UTIERIALES 23 spines, warm sepia; odor of acrid meal; latex salmon-color" (Zeller and Dodge 1937a: 633). Discussion.-Zeller and Dodge reported the fungus from Michigan on the basis of collections by Kauffman and Kanouse. GAUTIERIALES "Fructifications hypogeous, sessile; peridium wanting or present, when present stupose, loosely filamentous or pseudoparenchymatous; gleba gristly translucent and white when fresh, becoming brittle and brownish as spores mature; columella from a basal rhizomorph; basidia in a hymenium; septa usually gelatinous-cartilaginous, of gelified hyphae; basidiospores of various shapes, mostly broad fusiform, verrucose or longitudinally costate, brown" (Zeller, 1949: 41). GAUTERIACEAE Zeller recognized only one family, the Gautieriaceae, witi characters as given for the order. There is i genus, Gauticria. Gauticria Vittadini Fructification globular, pear-shaped or tuber-like, attached by a distinct basal rhizomorph; peridium variable in thickness, in some species fragile and disappearing by maturity, in others well developed, permanent, of 1 to 2 layers; gleba of anastomosed tramal plates, cavities labyrinthiform to cellular and lined with a palisade of basidia; columella simple to branched; sterile base usually present; spores colored, longitudinally ribbed. Gautieria morchelliformis Vittadini (P1. I, Fig. 3) "Fructifications spherical to oblong, 1-3 cm. in diameter, with a basal stalk-like, usually much-branched rhizomorph; columella rudimentary, merely a subspherical summit of the rhizomorph, from which rather thick septal plates radiate into the gleba; peridium thin in the early stages, soon evanescent; gleba ochraceous-tawny to hazel, cavities 1-6 mm. in diameter, sub

Page  24 24 PUFFBALLS AND) ALLIES IN MICHIGAN spherical to irregular, mostly radiating from the base; septa white when broken, stupose, about 75 p thick; basidia about as large as the spores, hyaline, granular, 2-3-spored, sterigmata filiform, as long as the spores; cystidia in the upper cavities of the fructification, not prominent; paraphyses clavate, septate, hyaline; spores 12-24 X 8-12.5 p, pedicellate, 1-2-guttulate, with 8-1o rather sinuous striae" (Dodge and Zeller, 1934: 699). Habit, habitat, and distribution.-Single or gregarious in soil, under hardwood trees and shrubs. It lhas been collected in southeastern Michigan from August to October. Like all members of this genus it is rare. l)iscussion. —Dodge and Zeller reported tlis species from New York, Oregon, and California. Gauticria graveolcns Vittadini "Fructifications spherical, 1-2 cm. in diameter, light ochraceous-buff to Prout's brown; stipe slender and fragile, up to 1 cm. long, 1 mm. thick; columella frequently reaching the center of the fructification, fruticose; odor very strong, as of decaying onions; peridium thin, composed of delicate, thinwalled, loosely woven hyphae, soon rupturing and disappearing; gleba ochraceous-tawny to cinnamon-brown, cavities spherical or elongated, minute, empty; septa 40-80 p thick, composed of small hyphae, compact; cystidia clayate to subfusiform, hyaline, often obscured by the spores; paraphyses linear, septate; basidia broadly clavate, 2-spored, 12-16 x8-9 p, with long, filiform sterigmata; spores ochraceous-tawny, apex rounded, mostly obovoid, pedicellate, 18-19X 11-12 p, often witl a large oil globule, usually with 7-10 prominent, smooth striae" (Dodge and Zeller, 1934: 695). Habit, habitat, and distribution.-Decply buried under leaf mold. At present the collections reported from Isle Royale, Keeenawi County, which were made by Brown and Povah, are the only ones known from Michigan. Discussion.-The ciaracteristic strong odor is an aid in recognizing this species.

Page  25 TREMELL OGASTRALES 25 TREMIELLOGASTRALES "Fructifications hypogeous or epigeous, mostly sessile; peridium of two or more layers, the outer of fundamental tissue, the inner of a gelatinous nature, continuous or interrupted by sutures of fundamental tissue; gleba centripetally developed, pulverulent at maturity; columella simple or wanting; spores spherical, echinulate, verrucose or cristate" (Zeller, 1949: 41). KEY TO FAMILIES OF TREMELLOGASTRALES i. Gelatinous inner layer of pcridium interrupted by sutures of fundamental tissue; spores echinulate to cristate................ TREMELLOGASTERACEAE i. Gelatinous inner layer of peridium continuous; spores verrucose............................................................ G ASTROS() ORIACEAE TREMELLOGASTERACEAE "Fructifications hypogeous or epigeous, mostly sessile; peridium of two or more layers, the outer of fundamental tissue, the inner of a gelatinous nature, interrupted by sutures of fundamental tissue; gleba centripetally developed, pulverulent at maturity; spores spherical, echinulate or cristate" (Zeller, 1949: 42). KEY TO GENERA OF TREMELLOGASTERACEAE i. Spores echinulate but not cristate; inner peridial layer thick and gelatinous, divided radially and periclinally into 2 more or less definite layers.................................................. Trem ellognstcr i. Spores echinulate-cristatc or merely cristate; inner pcridial layer thinner and not divided by sutures of fundamental tissue........... CClallhrogaslcr GASTROSPORIACEAE "Fructifications hypogeous, subglobose, from a single rhizomorph; peridium duplex, outer layer thin, of fundamental tissue (fibrous); inner layer gelatinous, continuous; columella simple; gleba pulverulent or deliquescent at maturity; spores spherical, minutely verrucose, slightly tinted or nearly hyaline" (Zeller, 1949: 42). Gastrosporium Zeller Zeller listed only i genus, Gastro orium, not yet recorded for North America.

Page  26 26 PUFFBALLS AND ALLIES IN MICHIGAN HYSTERANGIALES "Fructifications mostly hypogeous, globose or elongate, mostly from rhizomorphic strands; peridium simple or with 2 to 3 layers, or with an inner gelatinous layer (tramal peridium); gleba cartilaginous, gelatinous; basidia phalloid; spores smooth, ellipsoid to bacillar; tramal structure radiating from the base or as continuations from the mycelial strands, diverging from sterile base, or from gelatinous or cartilaginous columella. Rarely with a percurrent columella (Rhopalogaster)" (Zeller, 1949: 42). KEY TO FAMILIES OF HYSTERANGIALES i. Tramal peridium continuous, thick, gelatinous..............G GOELELLACEAE i. Tramal peridium not continuous..................................... 2 2. Tramal peridium thick, interrupted by thin plates of peridial tissue having unbroken connection with the fundamental peridium and sectors of the gleba.................................... PROTOI ALLACEAE 2. Tramal peridium thin and often poorly developed, interrupted by fertile or infertile cavities which are usually not filled by peridial tissue................................................ HYSTERAN(;IACEAE HYSTERANGIACEAE "Fructifications hypogeous, mostly with rhizomorphic strands or heavy mycelial spawn; peridium simple or with 2 or 3 layers; gleba cartilaginous, gelatinous, tramal structure radiating from the base or from a gelatinous columella (columella percurrent in Rho alogastcr); spores smooth, ellipsoid, or bacillar" (Zeller, 1949: 43)KEY '10 C.NERA OF HIYSTFRAN(;IACE.AE i. Fructification with a stalklike, unbranched percurrent columella...............................................................R h opalogaster 1. N ot as above........................................................ 2 2. Thicker branches of the columclla dividing the gleba into sharply delimited sectors; fructification with a prolonged, tapering sterile base............................................................. P iallogaster 2. N ot as above....................................................... 3 3. Fructification with short stalk, extending into the gleba, or a hemispherical, sterile base..............................................Jaczewskia 3. Fructification not stalked................................Hysterangium

Page  27 HYSTERANGIALES 27 Hystcraisgium Vittadini Fructification globular to pear-shaped or tuber-like, attached by radicating basally attached rhizomorphs; peridium 1- to 2 -layered, of woven hyphae or pseudoparenchyma usually partly gelatinized and in some species separating readily from gleba; gleba of tramal plates which usually gelatinize, anastomosed to form cavities lined with a palisade of basidia; columella simple or branched and arising from a sterile base; spores smootl or with a gelatinous exospore, hyaline or slightly tinted. Hysterangium clathroides Vittadini "Fructifications globose, becoming very irregular on drying, white to pale ochraceous buff or light ochraceous salmon when fresh, becoming buff-pink to onion-skin pink where bruised, drying ochraceous tawny to Prout's brown; fibrils variable from terete and free to innate or appressed; columella usuall y large and prominent; often branching near the base; peridiumi 220 -450 P thick, parenchymatous, the cells varying from 12 to 40 1 in diameter..,gleba green when fresh, becoming citrine drab or grayish olive to dark greenish olive on drying; cavities polyhedral to irregular, with a tendency to radiate from the columella, small, empty; septa 85-140 p thick, composed of large, thin-walled, loosely woven hyphae up to 5-7 P in diameter, finally becoming highly gelatinized; basidia long, irregularly cylindrical, 3-4-spored (mostly 3-spored); sterigmata usually short, although sometimes becoming i6-18 p long; spores acrogenous, olivaceous in mass, lanceolate, 12-19X6-8 p (averaging 15.3o 0.9 long), with a thick epispore which sometimes is slightly roughened and becomes loosened in age, sometimes papillate at apex, sometimes not" (Zeller and Dodge, 1929: (4 -95). Habit, habitat, and distribution.-Under needle beds among cedars in a swamp, but with deciduous trees nearby. Collected by Morten Lange (No. 1277) at the University of Michigan Biological Station near Douglas Lake, Cheboygan County, July 14, 1947.

Page  28 28 PUFFBALLS AND ALLIES IN MICHIGAN Discussion.-As the plant has been collected on many occasions in New York, its presence in northern Michigan is not surprising. Phallogaster Morgan Mycelium fibrous, much branched; pcridium of fructification consisting of 2 distinct layers, an inner and outer one, rupturing irregularly; gleba composed of numerous roundish irregular green masses or lobes attached to the inner surface of the upper part of the peridium; spores minute, oblong, hyaline. Phallogastcr saccatus Morgan (1'1. II, Fig. i) Fructification 2.5-5 cm. high, 1-2.5(3) cm. in diameter in widest part, obovoid to stipitate and subglobose; surface smooth at first, often superficially rimose, with broad shallow depressions, soon becoming visible around the enlarged part; color whitish tinged pinkish vinaceous or lilac; rupturing in a stellate manner or becoming perforated in the depressed areas, the manner of dehiscence rather irregular; interior at first having upper part filled by the gleba, the lower part (the stipe) stuffed by a white floccose material, both gelatinizing; gleba consisting of a number of green lobes or masses, at times quite distinct from each other and usually adhering to the inner surface of peridium as slimy masses, odor disagreeable at this stage; white to lilac rllizomorplhs conspicuous at the base or throughout the surrounding debris; spores sulbcylindric, 4-5 x 1.5-1.8 p, hyaline in KOH.1 Habit, habitat, and distribution.-Single, scattered, or in small clusters from single rhizomorphs. Throughout the state on very decayed d ood from late spring to early summer, or on into July in the northern part. It is considered rare, but once one knows wlhere to look for it, it can be collected several times during an average season. Discussion.-Tllis curious fungus, which closely resembles the stinkhorns in odor and spore characters, looks more like an I The solution used was 2.5 per cent potassium hydroxide.

Page  29 PHALLALES 29 ordinary puffball when first encountered. The foul odor does not develop until the gleba is mature and is never as strong as in the true stinkhorns (order Phallales). The field characters by which the fungus may be identified are the green chambered gleba, the pinkish to lilac color of the young fructifications, and the copious development of the white to pinkish rhizomorphs. Fitzpatrick (1913) studied its development. PROTOPHALLACEAE "Fructifications subglobose, hypogeous or epigeous; peridium usually thin, of primary tissue covering a thick gelatinous tramal peridium which is interrupted by radial sutures having unbroken connection with the peridium and gleba, which is gelatinous or cartilaginous, olivaceous or brownish, usually sectored by gelatinous plates radiating from the base or from a columella; cavities empty then filled with spores; basidial hymenium lining cavities; spores small, bacillar, olivaceous" (Zeller, 1949: 43). KEY TO GENERA OF IROTOPIIALLACEAE i. Gleba a powdery mass at maturity............................... C lvarula i. Gleba gelatinous-cartilaginous at maturity.......................Protubera GELOPELLACEAE "Fructifications subglobose, hypogeous; peridium thin, filamentous, surrounding a thick, continuous, gelatinous layer (tramal peridium); gleba dark, cartilaginous or gelatinous; cavities lined with basidial hymenium; columella simple or branched (pendant in G. hahashimnesis); spores small, smooth, colored" (Zeller, 1949: 44). Gellopellis Zeller There is "one genus, Gclopcllis, not known in North America" (Zeller, 1949: 44). PHALLALES Fructification in immature stage globose to ovoid; peridium of 2 to 3 layers, at maturity rupturing at apex to allow receptaculum to elongate; remains of "egg" present at base of receptaculurn as a volva; receptaculum variously shaped (stemlike in

Page  30 30 PUFFBALLS AND ALLIES IN MICHIGAN Michigan species); tissue pseudoparenchymatous, bearing the gleba which consists of an olivaceous slimy mass containing the small, bacillar, smooth spores. Zeller divided the order into 3 families: Claustulaceae, Phallaceae, and Clathraceae. Claustula is the only genus in the family of that name and is not known from North America. The genera he listed for the Phallaceae are: Xylophallus, Staheliomyces, AIutinus, Jansia, Floccomutinmus, Aporolphallus, Itajahya, Phallus, Echinophallus, and Dictyophora. In the Clathraceae he included the genera Clathrus, Colonnaria, Blunenavia, Aseroe, Simblun, Colus, Lateruea, Pscudocolus, Lysurus, and Kalchbre nn ra. I'HALLACEAE "Volva cupulate or sheathing, of 2 layers, the outer thin rind of filamentous primary tissue, the inner a thick, continuous, gelatinous layer; gleba surrounding the upper part of the receptacle; receptacle porous, stalk-like, with or without a bellshaped cap and sometimes with a continuous or meshy indusium; spores olivaceous or greenish, smooth, small, bacillar" (Zeller, 1949: 44). KEY TO MICIII(AN (FINTRA OF I'IIALLACEAE i. Rcccptaculum in the form of a stalk tapered to apex and with gleba covering upper part............................................ utinus i. Rcceptaculum with a pilcus.......................................... 2 2. Indusium (veil) rudimentary if present, not projecting below cap margin................................................................ P hallus 2. Indusium forming a distinct lattice-like skirt, extending for some distance below pileus......................................Dictyophora AIutinus Fries Fructification at maturity consisting of a basal volva (remains of the egg) and a stalk (receptaculum), which may or may not be perforated at apex and is of chambered structure; gleba slimy, olivaceous, foetid, and borne on apical part of stalk; no true pileus differentiated. Alutinus is easily distinguished from Phallus and Dictyophora by the characters used in tle key. In my experience the members of AIMtinus are the most common of the phalloids in the state.

Page  31 PHALLALES 31 KEY TO SPECIES OF AMutilnlS i. All parts except gleba white.........................M. caninus var. albus i. Stalk pinkish to orange red.......................................... 2 2. Stalk equal to near apex..........................M. caninus var. caninus 2. Stalk subfusiform or at least tapered from middle to apex........M. elegans Mutinus elegans (Mont.) E. Fischer Egg globose to ovoid, white with a pinkish tinge, attached by a white and often branched rhizomorph; fructification when expanded consisting of an elongated, hollow, spongy stalk 10-17 cm. long, tapering gradually upward to the blunt perforated apex, pinkish red, or paler to whitish downward; gleba borne on the upper part of stalk as a slimy, foul-smelling, olive-brown mass, structure of stalk beneath gleba same as elsewhere; pileus not differentiated, remains of egg forming a lobed volva around base of stalk. Spores 4-7 x 2-2.5 p. Habit, habitat, and distribution.-Around old woodpiles, on rich soil in gardens or lawns, on humus in the wood, or around very decayed hardwood logs and stumps. It fruits during wet warm weather in the summer and fall. Discussion.-I have seen it abundant around a very rotten willow log and stump at the Saginaw Forest, near Ann Arbor, Washtenaw County. This material compared best with that illustrated by Coker and Couch (PI. 5, 1928) under the name AM. Curtisii. The fungus is a curiosity and never fails to attract attention because of the pinkish red stalk. The buttons are often slow to open. I visited a group periodically for 2 weeks before they finally expanded. Mutinus caninus (Pers.) Fries Var. caninus (PI. II, Fig. 2) Egg white, 1o-15 mm. in diameter, 1-2 cm. long, attached by a rhizomorph; fructification a stalk 5-8 cm. high when expanded, 5-8 mm. thick, stalk equal to near apex where it is narrowed slightly to an obtuse usually perforated apex, red to orange-red or paler downward; base sheathed by a volva; gleba olivaceous brown, slightly odorous and covering the upper 2-3 cm. of the stalk with the exception of the very tip, the area of

Page  32 32 PUFFBALLS AND ALLIES IN AIICHIGAN the stalk under the gleba marked off from the sterile part by a difference in structure of thlc cells; spores 4-5 x 1.5-2 p. Habit, habitat, and distribuionl.-Gregarious to single on soil or very rotten wood in open grassy oak woods. The collection photographed was from the Edwin S. George Reserve near Pinckney, Livingston County, and appears to be typical M. caninlus. The funguts is apparently uncommon and fruits in late summer or fall. Discussion.-The difference in cellular structure of the stem of the sterile and fertile parts was not checked carefully on the material illustrated. The weak odor, sinall size, and typically eq(ual stalk are characteristic. In lMutins clegans the reccptaculum is tapered and a subfusoid structure results; compare Plate 5 in Coker and Couch (19)28) with tle illustration given here for AI. caninlus. A111tinlllS ca(inLnS var. albIus Zeller (1']. inI. Fig. i) Egg 1o-15 mm. in diameter, 10-20 lmm. long, o{void or finally obtusely conic, exterior white and unpolished; layer of gel 3-4 mm. thick; receptaculum 8-io cm. long, 8-11 mm. thick, equal, white, with honeycomb-like structure; the head 1-2 cm. long, 8-12 mm. thick, covered at first by a tough silvery gray membrane whichl gradually disappears exposing the olivaceous odorous gleba; apex of stalk not perforated; volva present as the persistent remains of the egg, usually with several lobes; spores 3.5-4.2 x 1.6-2 p. Habit, habitat, and distrib ttionl.-Clustered along an old dead, fallen birch tree, NIaple River, C b an County, July 13, 19.47. So far only known from the state by one collection (Smit1- 25702). Discussion. —The figure illustrates, in the specimen at the left, tlh very toutgh membrane covering the gleba. In the one next to it tile memblrane had disappeared and the gleba was starting to dri p off. I was unable to ascertain exactly what happened to tle lmemblran. )n tle specimen at the left in the illustration it was very to(ugll whIen I left the laboratory about

Page  33 PHALLALES 33 10 o'clock in the evening, but by the next morning it was gone. I have not observed such a tough membrane as this on either the typical variety or on Al. elegans. The claracter of the membrane with the difference in color suggests that var. albus slould, perhaps, be rated as an autonomous species. Phallus Persoon Fructification at maturity consisting of a volva, a stalk, and a pileus; pileus apical, outer surface smooth or reticulate; indusium (veil) poorly developed and not extending appreciably below cap margin; stalk hollow, of porous construction; gleba borne on tile outer surface of pileus, slimy, foetid. The name Ithyphallus is used by some authors for this genus, but this is not correct according to tle International Rules. There are 2 species in Michigan. KEY TO MICIII(AN SI'tLCIt.S OF Phallus i. Outer surface of cap distinctly reticulate......................P. itipudlicus i. Outer surface of cap granular.................................P. Ravclclii Phallus imnudicus Persoon Egg oblong to ovoid, whitish to pinkish, 3.5-6 x 3-4.5 cm., attached at base to rhizomorph; expanded fructification up to 25 cm. high, seated in tlie volvate remains of the egy; pileus 1.5-4 cm. long, outer surface deeply reticulated and tlhe depressions filled with remains of gleba (an olivaceous slime witlh a putrid odor), attached to the broadly expanded stalk; veil tliin, rudimentary, not perforated, not extending below cap margin; stipe 2.5-: cm. thick, tapered at each end, honeycombed, apex white; spores (3)3.7-4.2 x 1.3-2 p. Habit, habitat, and distriblltioln.-On tlhe ground usually around trees and shrubs. It apparently occurs late in the fall in Michligan, but seems to be rare. Discussion. — Mr. Walter Nickell, of the Cranbrook Institute of Science, hlas kodaclromnes of a plalloid he found near Bloomfield Hills, Oakland County, whlich unquestionably belongs to tlis species. Phallus impnudicus can be distingulished fromi

Page  34 34 PUFFBALLS AND ALLIES IN MICHIGAN P. Ravenelii at a glance by the reticulated outer surface of the pileus, and from Dictyophora duplicata by the absence of a veil below the cap margin. I have not seen fresh material which has been collected in Michigan, but the fungus has been reported from the state and is generally considered as common west of the Mississippi River. Phallus Ravenclii Berkeley and Curtis (I'l. IV) Egg typically large, 3-5 x 2-3.5 cm., ovate, whitisl, pinkish, or pinkish lilac, wrinkled at the base and furnished with a pinkish-lilac, well-developed rlizomorph, or several eggs along a single rhizomorph, consistency gelatinous-pliant (because of gelatinous inner layer); expanded fructification with a stalk ioi6 cm. high, slightly tapered upward to nearly equal, 1.5-3 cm. thick at base, base enclosed in the remains of the egg which forms a volva, surface yellowish fading to white, honeycombed; pileus, attached around a raised white disc at apex of stipe (disc perforated at center), in the form of a thin membrane which is granular rather than rugose or reticulated; gleba evenly distributed over pileus and olivaceous gray; veil membranous and white but not ordinarily extending below the edge of the pileus; spores 3-3.5 x 1.5~. Habit, habitat, and distribution.-Densely gregarious, rarely single, around or on very rotten logs or other ligneous debris, during the summer and fall. This fungus has been recorded for the state in late August, but only in very wet summers. It is more abundant after the fall rains if the weather remains warm, and is not uncommon in at least the southern part of the state. Discussion.-The granular surface of the pileus together with the color of the eggs and mycelium amply characterize the species in the field. Dictyophora Desvaux Fructification, when expanded, consisting of a pileus, an indusium, a stalk (receptaculum), and a basal volva (remains of the egg); pileus campanulate, reticulated, attached at apex of

Page  35 PHALLALES 35 stipe; indusium a latticed pscudoparencliymatous pendant membrane attached to apex of stipe beneath pileus, hanging free from stipe and projecting some distance below cap margin; spore mass olivaceous, slimy, fetid, covering reticulate surface of cap; stalk typically large, latticed, perforate at apex. The indusium (veil) and presence of a distinct pileus are the diagnostic characters. There is only i species in the United States. Cunningham reported 2 species, D. indusiata and D. multicolor, from Australia. Dictyophora duplicata (Bosc.) E. Fischer (11. III, Fig. 2; I'1. V) Egg globular to flattened, 4-6 x 4-6 x 5-7 cm., white to whitish or tinged flesh color to brownisll on exposed surface, sometimes plicate around the attachment of the well-developed white rhizomorph, sometimes with more than 1 rhizomorph, consistency pliant due to the inner gelatinous layer, finally rupturing by an irregular slit; expanded fructification consisting of a fleshy-gelatinous volva (remains of egg), an elongated stipe, a pileus and an indusium; pileus a thin membrane attached to apex of stipe and pendant down the sides, upper surface reticulate and bearing the gleba, under surface smooth and wlite; indusium attached under the cap at apex of stipe, lianging 3-6 cm. or more below lower margin of pileus, netlike in pattern, the meshes about 1-2 mm. broad or smaller near margin, white to pinkish; stipe 3-6 cm. thick at base, narrowed upward, apex perforated, white, chamibered; spores 3.5-4 x 1.5-2 [, smooth. Habit, habitat, and distribt l/iol. —Single or in groutps in tlie woods, often near uprooted trees, around tlhe base of dead trees, or on humus. This fungus fruits in the late summer and fall and is frequent in tlie thin sandy oak woods in the southern part of the state during warm wet weather. Discussion.-Once while collecting fungi witlh other students we smelled a plhalloid and tried to trace it down in order to learn the species, but it seemed to keep moving. Finally, we noticed that an old man was also in the woods collecting fungi, and as I worked over toward him I realized that a phalloid of

Page  36 36 PUFFBALLS AND ALLIES IN MICHIGAN some kind was very close. After greeting the old gentleman we engaged in conversation, and a few minutes later he took off his cap and slhowed us a large fruiting body of D. duplicata in it. After some exclamations on my part he informed me that he used it in this manner to cure his rheumatism. The species has no properties to effect such a cure. The fungus is at once distinguisled by the reticulate upper surface of the pileus and conspicuous veil, lwich is nearly always badly torn as the stalk elongates. LYCOPERDALES "Fructifications mostly epigeous, sessile, single or in groups on a stromatic layer, rarely substipitate, globose, pyriform, etc.; peridium 2-4 layered, deliiscing by an apical pore, by several pores, by irregular or stellate cleavages, or crumbling at maturity; gleba wholly fertile or sterile below, becoming a powdery mass or chambers breaking apart and forming at maturity small hollow peridioles; basidia borne in a hymenium; capillitium present (except in Arachniaccac)" (Zeller, 1949: 47). This is the largest order of Gasteromycetes and the one most fully represented in the Michigan flora. The combination of spore powder and capillitium to form the gleba is typically diagnostic. KEY TO FAMILIES OF LYCOPERDALES i. Fructification brittle and disintegrating at maturity; glebal chambers each with its surrounding wall separating to forii small sandlike particles (peridioles)....................................ARACIINIACLAE i. Fructification persistent; glebal chambers disintegrating into a powdery mlass......................................... 2 2. Exoperidiutm opening stellately at maturity (cf. Ay)cclasirS1u1)... GEASTRACEAE 2. N ot as abov........................................................ 3 3. Peridium 2- to 3-layered, indehisccnt or rupturing irregularly at apex; capillitium unbranclhed............................... MESOIIEILIACEAE N~. N ot as above........................................................ 4 4. 'lhreads of capillitium witl spinelike side branches........ MYCLNASTRACLAE 4. Threads of capillitium smooth, often variously branched but not as alove............................................................ 5 5. Fructifications single to ilany on a stroll a....................BROOMEIACEAE 5. Fructifications not on a stiomaL.............................LYCOPERDACEAE

Page  37 LYCOPERDALES 37 ARACIINIACEAE "Fructiftifications epigeous, small; pcridium thin, fragile, breaking irregularly or crumbling at maturity to liberate the peridioles; gleba made up of numerous spherical chambers lined with a hymenium, forming at maturity a mass of minute, separate peridioles which are like grains of sand; capillitium and sterile base none; spores smooth" (Zeller, 1949: 47). KEY '0 (,LNiRA OF ARA\.(:IIN.\:ACI.AE i. Fructifications sessile, columella wanting........................ raiclh ion i. Fructifications stipitate, columella present........................ Arncosa Arachnion Sclwieinitz Fructification globose to subglobose; peridium very fragile; gleba a mass of small particles (peridioles) which contain the spores; subgleba (sterile base) none. The characters of the genus are also tlose of tlhe family Araclhniaceae, to which tlis curious fungus belongs. Altthough of simple organization, tile genus is interesting in tlat it has separate peridioles as do members of the Nidulariales, an order in which this type of structure reacles a high development. Aracliliou (alblmin Schweinitz Fructification 0.5-1.5 cm. in diameter, globose to subglobose but with a base tapered to point of attachment, attached by a small but rather persistent rootlike rhizomorph; surface pure white when immature, in ae, yellowish, smooth, cleveloping an areolate pattern in drying; wall of spore case very tlin and fragile, breaking up into fragments at maturity; gleba white tlhen becominig grayish to brownisll, brownis olive, or dark dral), composed of a large number of minute empty chambers lined by tlhe hlymenium, context betwem-en thle chambcrs made up of loose delicate interwoven hyphlae which break down at maturity freeing tlhe clambers as small sandlike particles (sometimes referred to as peridioles); sterile base none. Spores short-ellipsoid, sulglolose or a few comlpressed so as to be wider than long, 4.5-4.4 x 3.5-4.2 p (4.2 x4.5!u) smooth,

Page  38 38 PUFFBALLS AND ALLIES IN MICHIGAN thick-walled, witl a basal stump of a pedicel in most, rusty brown in iodine. Halbit, habitat, and distribltion.-The only Michigan collections I have seen were made on sandy soil in a pasture near Itlaca, Cratiot County, by Vi(tor Potter during the seasons of 1947 and 1948. The species is rare, or rarely collected, in Michigan. Disc(ssionS. — hTlcn the wall of the spore case breaks, the granular mass which is the gleba is easily dispersed and soon no sign of tlie frtuctification remains. The immature specimens are easily mistaken for tlhe lltt.)on stages of otlher puffballs. Consequently, tlle average collector is likely to pass it by unless he is making a special scarcl for it. LYCOPERDACEAE "Fructifications single or in groups, mostly epigeous, subglobose to pyriform or nearly stipitatc; gleba wholly fertile, or sterile below; outer peridium mostly a layer of pseudoparenchlyma, rarely with a rind tlat is skin-like or permeated witl soil particles, wholly or partially disintegrating at maturity, laying tlhe inner periditum bare; inner peridium usually papery and thin, rarely corky and tlick, usually delhiscing by an apical pore; rarely (in Lycopcrdopsis) tlle two layers adhere to each otlier forming a simple pseudoparenchymatous rind, usually opening by an apical pore; capillitium well developed, sometilmes falling into pieces" (Zeller, 1949: 48). KEY '1OC (;I:NRA OF LYC()'OPRD)ACLAE i. PI'rid(iuln lmade uI) of a looscly interwoven endoperidium and a closely ad(llrillg p)SCidi(ll)ilarlnci in1llaltous Col)eri(iidum..............Lycolpcrdop)sis I. P'lridilllll consisting of a nlCllllranous endoperidiulm which is laid bare at maturity by sloughing oft of exoperidium......................... 2 E. Endloperidiull ope()ing by an ap)ical lpore (sometimes basal in Disciseda). 3 2. Endo)eri(iuml dehliscing by irregular rupture......................... 6 3. (Capillitilim threads 1much blranclhed from a main stem, branches slender anid with tapered pointed ends; spores with long pedicels........Bovistella 3. Tlhlreads of capilliti umi withonut a conspicuous main stem and without a distinct systel (of branl ches......................................... 4 4. Exoperidiln separating lin endol)cridium circumscissilely exposing about half or more ol endo:peridium (fructification then resembling an acorn in its culp)........................................... D isciseda

Page  39 LYCOPERDALES 39 4. N ot as above....................................................... 5 5. Capillitium accompanied by membranes at maturity........... Morganella 5. Capillitium not as above..................................... Lycoperdon 6. Capillitium much branched (main stem with branches tapered to points, the threads free from the inner peridium and base.................. 7 6. Threads of capillitium smooth or granular, lacking a conspicuous main stem or conspicuous branching.................................... 8 7. Capillitium threads not thickly interwoven, dispersing readily when exposed.......................................................Bovista 7. Capillitium threads densely interwoven into balls..................Lnopila 8. Capillitium threads simple, smooth, short, with sharp ends...... Bovistoides 8. Capillitium of long threads or these breaking up easily, sparingly branched......................................................... 9 9. Endoperidium breaking up and falling away, usually leaving a sterile base covered by a definite membrane; capillitium threads usually breaking up..................................C............ Calvatia 9. Endoperidium cartilaginous, very thin above, splitting into several irregular toothlike segments at apex........................Arachnio sis Calvatia Fries Fructification typically large, subglobose to pyriform, in most species with a well-developed sterile base; outer peridial layer thin and smooth to granulose or consisting of warts or spines, inner peridial layer thin and fragile and at maturity fragmenting from apex outward to expose the gleba; gleba lilac to yellowish olivaceous depending on species, typically white when young; sterile base either fibrous or chambered, poorly developed in a few species, in many persistent long after spores have been dispersed; capillitium of long tlreads, branched to sparsely branched, septate to continuous, in some species fragile and breaking into short segments, attached originally to inner wall of spore case; spores typically globose and possessing a thin hyaline envelope into which minute spines may project. The genus is best distinguished from Lycoperdon, with which it intergrades, by the manner in which the thin, inner wall fragments to expose the gleba. In Lycoperdon this inner wall is less fragile and the spores are liberated through an apical pore or slit. KEY TO SPECIES OF Calvatia i. Gleba distinctly purplish at maturity................................. 2 1. Gleba olivaceous to dark yellowish brown............................. 3 2. Sterile base prominent and with distinct chambers........... C. cyattiiformis

Page  40 40 PUFFBALLS AND ALLIES IN MICHIGAN 2. Sterile base poorly developed, chambers indistinct................ C. fragilis 3. Fructification large (1o-45 cm.) globose or nearly so; sterile base very inconspicuous if present..................................... C. gigantea 3. Fructification with a distinct, chambered, sterile base................. 4 4. Fructification at first covered with a coating of soft spines which may be almost granular in texture and break down to a powder, sterile base much elongated................................C. saccata var. elata 4. Surface of fructification not as in above choice........................ 5 5. Capillitium with large nearly round pits....................C. craniformis 5. Capillitium with elongated to linear pits........................C. Bovista Calvatia cyathiformis (Bosc.) Morgan (P1. VI, Fig. i) Fructification 6-15 cm. thick at broadest point, 9-20 cm. high, varying from globose to depressed-globose above a pinchedoff base to broadly pear-slaped, at times nearly turbinate, often sulcate to deeply wrinkled from the base upward to broadest dimension; outer surface smooth at first but often becoming areolate over enlarged upper part and in some developing characteristic flat scales; endoperidium thin and delicate, slowly scaling away together with exoperidium at maturity to expose the deep purple-drab gleba; gleba white at first, changing through yellow to deep purple-drab; sterile base (subgleba) well developed and persistent, chambered, remaining as a cuplike structure long after spores have been sled. Spores globose, 4.5-6.2 p, with a thin, hyaline, gelatinous envelope through which the numerous distinct echinulations project, dark yellowish brown in iodine; capillitium of threads up to 5 p in diameter, with numerous septa and tending to break up at the septa, sparingly branched, the walls slightly thickened and with numerous small pits, pale yellowish brown in iodine. Habit, habitat, and distribution.-Solitary, scattered, or in "fairy rings" in pastures and grassy fields. In Michigan this species fruits abundantly during the wet spells of late summer and fall. It occurs throughout the state and is the most common of the larger species of Calvatia. Discussion.-Coker and Couch reported it as excellent food, if young specimens wlhich are wllitc clear through are fried like an egg plant. Shantz and Piemeisel (1917) in their study of

Page  41 LYCOPERDALES 41 this fungus in Colorado estimated the age of some fairy rings of it as 420 years, that is, as old or older than many large forest trees. Few people ever think of a fungus mycelium as very longlived. In Colorado it was noted that the rings advanced about 24 cm. a year. In fairy rings of this species the grass is stimulated just behind the ring of fruiting bodies. In rings of some other fungi the grass is dead in this zone. Calvatia cyathiformis can be distinguished when mature by the distinctly purplish gleba, the areolate pattern of the surface of the upper enlarged part, and the well-developed sterile base. It is often attached to the soil by a thick rhizomorph with much included debris. Calvatia fragilis (Vitt.) Morgan (I'1. VI, Fig. 2) Fructification depressed-globose to obovate, 3-7 cm. in diameter, 2-4 cm. high; base tapered to an obtuse point of attachment, radially wrinkled to plicate from base upward in some specimens, with an outer coating of whitish thin matted felt which flakes off in thin sheets to expose the thin, very fragile purplish inner peridial layer, the inner layer breaking up into flat irregular areas which fall away to expose the purple gleba; gleba cottony but persistent and not easily broken up (as it is in C. Bovista); subgleba (sterile base) present as a thin layer in the constricted part of the fructification, chambers indistinct. Spores purplish in mass, globose, distinctly warted, apedicellate, 4-5, in diameter, without the echinulations, 5-6.3 p including the projections, echinulations in the form of hyaline rods; capillitium of relatively short segments, cross walls numerous, walls slightly thickened, pale but tinged under the microscope, 3-6 p in diameter, minutely pitted. Habit, habitat, and distribution.-Two fructifications were collected in a sandy meadow near Riverdale, Gratiot County, by Victor Potter, October 13, 1948. Discussion.-The specimens collected by Potter are intermediate between C. cyathiformis and C. fragilis, in that the spores are spiny as in the former, but in all other characters arc

Page  42 42 PUFIFBALLS AND ALLIES IN MIICHIGAN like the latter. I have exaimined abundant material of C. fragilis from Texas, determined by the late Dr. Zeller. C. fragilis, like Lyco/cr(ldo pu/lchIrrim7111n, appears to be distinctly southern in its range and very common in central Texas, so that Potter's collection and its variation in spore character are particularly interesting. Calvatia giganteac (Pers.) Lloyd (P1. V\II) Frulctification subglobose to globose, or in large ones depressed-globose, (10)20-45 cm. wide and 20-35 cml. tlick, attached to tlle ground by a strong cordlike basal rhizomorph, white to creamy white when * young: surface smooth to finely tomentose and with tle feel of chamois: outer peridial layer of wall thin and breaking up into areolate patches which shrivel, becomie brownish, and may disappear: inner peridial layer of wall finally breaking up into irregular fragments and falling away in pieces exposing the gleba: gleba white when young but beconming yellowisl as spores mature, soon olive-yellow and when old rusty oclhraceous' subglleba very shallow or practically obsolete, not chambered. Spores 3.5 —4.5 P, globose, apiculate, with a hyaline envelope into whiclh minute echinulations project; capillitium of sparingly branched septate tlreads 5-6.5 1 in diameter, olive-yellow in O()H and withl slightly tlickened walls furnished with minute pits. Hallit, habitiat, and distril)utioon.-Single to gregarious on low ricl soil in b)rtslhy places, low woods, pastures, or the like. In soutlheastern Michligan it usually fruits about the middle of September, if the weather is favorable. Calvatia gigantea is one of tile common large puffballs of the state. The best fruitings I lhave seen were in and along drainage ditches in the lowlands east and northeast of Soutl Lyon, Oakland County. I)isclssionl.-Thlis is tile species usually sold on farmers' markets in soutlhern Mlichigan and the one most people think of whlen edible puffballs are mentioned. Thle large size, tlie whlite nearly smootl outer surface when young, and tlhe lack of any ap

Page  43 L YCOPERDA LES *1. precial)le sterile blase distinguislh it. llhe color of tile gleba is distinctive also, but people collecting for tlie table are not interested in specimens which slow tllis character. To be good to eat thle ptffball nmust be white clear through. Any tinge of olive or yellow is a sign of approaching maturity. ()ne cannot judge degree of maturity by size alone, for small balls io cm. (or even less) in diamneter may b)e more mature than other ttwice as large. In bu)ying puffblalls on tlhe market one should be sure to have tlhem cut in lalf. lWhen this is done one can examine tlhem for worm lholes, which appear as minute pinlioles in the flesh. If they are wormy, tile hloles will be most abundant near tlle point of attachment to tlhe rhizomorph. If the specimens are perfectly whlite witlin and otlerwise firm, any wor)my parts can simply be trimmed off. Buller (1922, p. 13 9) estimated that a spore case 40 x 20 x 20 cm. contains over 7 trillion spores. From tills one can scarcely imagine the number of spores slled by the hlundreds of tllese puffballs whicl develop in tlhe fall during a good season in just one of thle southern counties. Some monstrous specimens tip to 5 feet in tlhe widest dimension have been recorded in tlle literature. Calvatia craliformis (Schw.) Fries (1'1. VIII) Fructification 6-15 cm. at widest part, up to 20 cm. higl0h, obovoid, or top-shaped to elongated pear-shaped, upper part often deeply wrinkled, with a well-developed and often thick sterile base tapering downward to soil, where it is 2-6 cm. broad and attached by rhizomorphs: surface covered by a very thin, papery, smooth, unpolished to minutely furfulraceous outer layer which is pale avellaneous to dulll tan and scales off at maturity exposing the inner peridial layer, whiclh is also tlin and fragile and some shade of yellowish brown at Imaturity, tile upper part cracking into small patches and falling off exposing the gleba; gleba white tlhen yellowisl and sordid olivaceous, near Isabella color (Ridgway), eventually darker and more brownish, near Saccardo's tumber (Ridgway); stublgleba occupy

Page  44 441 1PUFFBALLS AND ALLIES IN MICHIGAN ing stalklike 1)ase, struc11(ti-re rat 1t' ' sp)onlgy l)bcatuse (' o la'rg clhainbers, persisting after tlle spores lhae been dispersed. Spores glolose, 2.5-3.5 p, with a tlin hyaline envelope into which a few ninute echinulations project (appearing smooth under ordinary [NA. 1.25] oil-ilmmcersion lens), with a broken stump of a pedicel or apedicellate; capillitium of threads with few septa, 2-6 p in diameter; lwalls thickened slightly and with large circular pits 1-3 p in diameter, often breaking at the pits. IHa, it 7ai, haitat ld distriblio.. —Single to scattered in open oak woods. As far as my records go Calvalia craliformis fruits later than C. cyathiformis and C. (i(rgaitca, which usually means October in soutlern Mlichigan. It appears to be rather rare. I)isciissiol.-According to Cok-er and Couchl this flngus is readily distinguished from C. Bovista by the circular instead of linear pits in tlhe capillitium and by the diameter of the threads being about tlat of the spores, rather than twice that of the spores (7-11 [17] p[) as in C. Bovista. On the basis of this distinction my collections all belong in C. craltiijoiris, though others have reptortcd C. Bo-isla (under the name C. caelala) as frequently collected in Michigan. I have found no specimen amnong cither Kauffman's or Swartz's collections which could be placed in C. Bovista, if emphasis is placed on the characters of the capillitium. Tle fructifications of C. craniforiis are sometimes so wrinkled and folded as to give tlem the appearance of a brain, but the character is not constant. Calvalia Bovista (Pers.) Kambly and Lee "Plant large to very large, sterile base ample, either long and cylindrical or slorter and tapering to the pointed base; cortex a flocculent layer lwhich is thicker above and there usually areolated into more or less prominent warts, thinner downward, but witll a tendency tlrotughout to form more or less stellate, flattish areas; color wlhitislh, then pallid yellowish to brownish. Inner peridiui tlhin, breaking up into fragments at the top, then downward, exposing tlhe bright olivaceous gold to brownish olivaccous gleba, wliclh is very fragile and powdery. Sterile base

Page  45 LYCOPERDALES 45 persistent, up to!9.5 (:1. tlick aiid ol cm1. highl, fulrrowed below and pinched to a point; tlhe cliambers of the subgleba distinct, empty and moderately large, extending up the sides for several centimeters as a tapering margin to the cup. "Spores (of the plant from Pawling, New York) spherical, smooth, with a short mucro and distinct oil drop, 3.8-4.5 p. Capillitium threads up to 17 p thick, usually 7-11 p, somewhat branched, easily breaking up at maturity; walls with narrow (about 1 p wide), linear, sinuous pits." Discussion.-The description given is quoted from the account of C. aclata by Coker and Couch (p. 68). This species, which grows in fields, lawns, pastures, or along roads in the late summer and fall has been frequently reported from Michigan under the name C. caclata, but I can find only a single collection in the University Herbarium, one made at Whitmore Lake by Mains and identified by Kauffman, which has the capillitium of this fungus rather than of C. craiiiformis. This collection, however, lacks the characteristic sterile base typical of C. Bovista and in shape and color of gleba is almost identical with small specimens of C. gigantea. The other collections identified by Kauffman and Swartz all belong to C. craniformiis. Good material of C. Bovista is in the herbarium of Michigan State College, identified by Barnett, but with no data as to locality or time of collection. The capillitium and spores are typical but some of the fruiting bodies have a well-developed pseudorliza. A collection by Clarence H. Kennedy from the Batchlawana Bay area in Ontario was made in 1949 and is in the University Herbarium. Both the concept of this species and its name have had a varied history, but if the Internatiolnal Rules are followed, the species epithet Bovista must be used and I have, therefore, accepted the combination made by Kambly and Lee. Calvatia saccata var. elata (Massee) Hollos (P11. IX) Fructification resembling that of Lycoperdon, but with a long sterile base; head 3-6 cm. broad, over-all length 8-12 cm., elon

Page  46 46 PUFFBALLS AND ALLIES IN MICHIGAN gate-pyriform or subcapitate on a stipe whlich enlarges upward, white when young and enlarged part covered with a granular to soft, floccose coating resembling that of Lycoperdon unmbrinumr var. floccosum, stipitate part with a faint powdery coating to nearly glabrous; wall of spore case very thin and papery, becoming pale brown in maturing, and then breaking up into flakes which eventually fall off exposing the gleba; gleba white at first, changing through yellow to olivaceous and finally olivaceous brown to dark yellowish brown; sterile base (subgleba) occupying the long, stemlike base, chambered. Spores globose 3.7-5 P, with a thin hyaline envelope into which project numerous echinulations, usually with a short stump of a pedicel; capillitium threads long, sometimes branched, 4-6 p in diameter, walls thickened and with numerous small pits on main branches, branches tapered to acute apices, rusty brown in iodine. Habit, habitat, alnd distribution.-Single or 2 to 3 near each other on humus in oak woods. I have 3 collections from near Milford, Oakland County, of immature to scarcely mature specimens, made in late September or October. Specimens which apparently had overwintered are among Kauffman's collections in the University Herbarium. They were obtained in May. The fungus seems to be rare in Michigan. Discussion.-Tlis fungus can best be characterized as having the appearance of a large member of the genus Lycoperdon, but is one in which the wall of the spore case breaks up as in Calvatia. It is most likely to be confused with Lycoperdon umbrin umn var. floccosum, especially in the immature stages. In the literature it is reported either as C. elata (Massee) Morgan or as C. saccata var. elata (Massee) Hollos. Even those who give it the rank of a species usually state in their discussion that it is only the American form of C. saccala. Coker and Couch, who ranked it as a species, emphasized the difference in spore size (3.5-4.4 p as compared to 4.5-5.5 p). My measurements (which include the spines) are intermediate.

Page  47 LYCOPERDALES 47 Lycoperdon Persoon Fructification small- to medium-sized, i-io cm., globose to subglobose or pear-shaped, some with a prominent stemlike base, attached to substratum by one or more rhizomorphs, consisting of a surrounding wall which encases a mass of spores intermingled with threads; wall composed of 2 layers known, respectively, as the exoperidium (outer wall) and endoperidium (inner wall); outer wall consisting of a spiny, warty, granular or nearly smooth floccose coating, which ordinarily sloughs off at maturity; inner wall forming the membranous more or less rigid spore case and giving shape to the fructification, its consistency at maturity somewhat membranous or papery and its surface smooth or with characteristic markings, typically opening at apex by a pore (often called a stoma) through which the spores escape; gleba (the combined mass of spores and threads in the fruiting body above the sterile base) colored and with typically thick-walled colored threads, collectively called the capillitium, which originate from the inner wall or adjacent sterile tissue under the gleba; subgleba, a sterile tissue chambered much like a honeycomb (in most species) but with smaller cavities, occupies the basal part of the fructification or the stemlike base if one is present; spores globose to subglobose, rarely ellipsoid, with or without a pedicel, typically surrounded by a thin hyaline envelope into which project minute echinulations (needle-like projections) or warts, the colored wall thick and usually the only one seen unless a good high-power oil-immersion lens is used, spores smooth in a few species. KEY TO SPECIES OF Lycoperdon i. Sterile base not chambered, greatly reduced in size, or absent; if slightly chambered the cavities less than 0.5 mm. in diameter............... 2 1. Sterile base typically well developed................................. 6 2. Surface of fructification conspicuously pitted at maturity; spiny coating cinnamon buff to brown; habitat on mossy logs..... L. subincarnaluml 2. Surface at most obscurely pitted or reticulate; typically smooth or with a slight furfuraceous covering..................................... 3 3. Surface with inconspicuous paler spots vAhen spiny coating has just sloughed off; spiny coating white when immature; habitat in grassy places; capillitium hyaline in KOH..........................L. Curtisii

Page  48 48 PUFFBALLS ANI) ALLIES IN AIICHIGAN 3. Not wiltl ab(,ove comlninaltionol (,1 c( l.r l ct( rs............................ 4 4. Fructification a Irilliant golden oranlge-y)ellow lwhien ilmatlll;lti'.L.. colo)oralu 4. Fructification white or whitish young............................... 5. No sterile base present; fructification small (1-2 cm.) spores globose or nearly so............................................... L. i)usil7um 5. As above but sporcs ellipsoid...........................L. oblongisporu)i 5. Sterile base typically presenlt; fru iting bodies larger (3.5 c1.)............................................................... L. p oly) or m 1 11 6. Spiny coating soon blocked out into warts with sharp tips and separating from the surface in chunks or siceets.....................1... mnrgtinatum 6. Not as in above choice............................................. 7. Typically growing on decaved hardw(ood logs, stum)ps. or delris; remains of outer coating more or less p)crsistent and rough to the touch as is a fine file; spores smnoothL..............................L. pyrifor1 ec 7. N ot as above................................................ 8 8. Gleba typically olivaceous blrown to dark brown but not tinged pturplish. 9 8. Gleba in fully mature p)lants tinged purpi)lish................ 13 ). Habitat on mosses ('o!virlr/ /llil S. c S',Il I),,, etc.)............7L. liscor/iN)7 ). H abitat not as above............................................... o 1o. Spiny coating tinged lavender on ilmmature specimcns; spines split at base iinto 2 to 4 parts but tips ulnited............................ Prckii lo. Not as in albove choice............................................ 11 ii. Spines cone-shaped and leaving distinct spots on the surlitcc when they fall off........................................... L. p)erlatnu 11. Spiny or warty layer variablle b t l no(t as above...................... 12 12. Spores sutbglobose to bIroadly oval. with long persistent pecicels............................................................. L. p ed icellatu m 12. Spores glol)ose, Iped(icels molstly broken oil and floating free in mounts........................................................... L. u 7 ) rinum1 a Outer coatinlg sctlrfy to gran.u!lar...................... ar. n11ibri7nu b Outer coating of long flocculent spines.................var. floccosuin 13. Outer coating smooth, finally sloughing off in form of thin plates............................................................... L,. ^ri z laium 13. Spiny covering of granules interlnixed with longer hairlike spicules............................... L i. I.brinumi (especially var. atropllrpr11 i m) 13. ()uter coating of Iong spines (2-6 111.)............................. 14 14. Surface smooth when spines have fallen off...............L. pulcherrimumn i4. Surface reticulated at least for a time after spines have fallen off...... c i......................................................... L. c Ii n t Lycopcrdon si) 11 icarnat um Peck (PI'1. X, Fig. i) Fructification 1-3 cm. broad at widest part, short-pyriform, globose or depressed globose, attached to substratum by one or several whlite rilizomorphs, color cinnamon-buff to cinnamonbrown when youlng, and covered with minute, separate, brown

Page  49 LYCOPERDALES 49 spines and warts in groups with their tips united and connected by their bases with brown tissue, the scurfy coating gradually wearing off and at full maturity the surface conspicuously pitted and much paler (pallid to pale tan), the pitting developing at or after maturity and evident even though spiny layer does not slouglh off; gleba with distinctly radiating tramal plates which in immature specimens are reminiscent of Geastrum, in mature specimens grayish or tinged purplish; no columella observed in material I have examined; sterile base rudimentary to wellformed depending on whether the fructification is pear-shaped or globose. Spores globose, pale brownish in KOH, rusty brown in iodine, witlI the usual tlin hyaline envelope into which project short spines or plugs, 3.5-4 p in diameter, apedicellate but some broken pedicels present in mounts; capillitium of very pale threads, hyaline to slightly yellowish in both iodine and KOH, witl relatively thin walls and much incrusting debris, sparingly branched, frequently septate, 4-7 p in diameter. Habit, habitat, and distrib ltion.-Gregarious to scattered on moist mossy logs. It fruits during the late summer and fall, but has rarely been collected in the state. It may be more frequent in the northern areas. Kauffman had 1 collection, made near New Richmond, Allegan County, in 1911. Discutssion.-My description has been based largely on a collection by Kauffman from Mount Gretna, Pennsylvania, September 7, 1924, and one by Krieger at Rhodes Island, Magnetawan, Ontario, August 17, 1922 (Kelley-1414). The following notation is on the label of Krieger's collection: "This species caused diarrhea in L.C.C.K. and Dr. H. A. Kelley." In view of this information any one finding the fungus in sufficient quantity to eat slould be careful. As can be seen at once from the illustrations, in the young stages it is very easily distinguished from L. pyrifornmc by tlle type of scales and by tlhe pits which finally cover the surface. The 2 species are also readily separated under tile microscope by tlhe characters of tlhe capillitium.

Page  50 5~ PUFFBALLS AND ALLIES IN MICHIGAN Lycoperdon Curtisii Berkeley Fructification small, typically 1-2 cm. broad, globose to subglobose when young but often misshapen from mutual pressure, sessile and without conspicuous rhizomorphs; surface at first covered by crowded spines united by their tips into stellate groups, fine granular material between the spines, spines falling off tardily and over small areas around apex first but finally over all of upper or exposed surface of spore case, the granular material remaining longer as a powdery coating, the pale-brown surface finally becoming smooth or nearly so; gleba olivaceous and more powdery than in most species (because threads of capillitium are not as rigid); sterile base very poorly developed, occupying only the region immediately adjacent to attachment of fructification to substratum, chambers small, at maturity purplish brown. Spores globose, small, 3-3.5 p in diameter, with a thin hyaline envelope into which echinulations project, olivaceous in KOH, rusty brown in iodine, with a large central oil drop; capillitium of hyaline (in KOH) to yellowish (in iodine) thin-walled, septate hyphae, 3-7 P in diameter, with much debris adhering to them. Habit, habitat, and distribution.-Usually densely cespitose to gregarious in pastures in the fall. It is to be expected in grassy areas, such as golf courses, lawns, abandoned fields, and vacant lots. Most of the material I have seen in the southeastern part of the state was observed between September 30 and October 15. Althouglh not as common as L. marginatum, when it does appear, it is likely to occur in larger quantities. It is a common species in the Lower Peninsula. Discussion.-I have here followed the concept of Coker and Couch and considered L. WIrightii Berkeley to be a synonym. The capillitium of this fungus is very similar to that of L. subincarnatum, but the 2 species have little else in common. Although immature stages of L.marginatum and L. Curtisii are similar in appearance, there is no good reason why anyone equipped with a microscope should confuse them. The echinulate spores and pale capillitium readily distinguish L. Curtisii.

Page  51 LYCOPERDALES 51 Lycoperdon coloratum Peck sensu Coker and Couch Fructification globose to subglobose, (1.5)2-4 cm. in diameter, pinched off at base and often sulcate around point of attachment, attached by one or more rlhizomorphs, some specimens with a rootlike process made up of dirt held together by rhizomorphs; surface of spore case brilliant golden orange-yellow until maturity when it changes to bronze and finally to dull brown, covered with minute harsh discrete warts or nodules which slowly become dark brown to nearly black, surface finally more or less glabrous at least around the pore but warty coating very persistent; gleba white when young, changing through yellowish to yellowish olive and at last olive brown to coffee brown; sterile base not well developed and chambers inconspicuous (best seen under a lens right above point of attachment to substratum). Spores 3-4.2 p in diameter, globose, with a thin hyaline envelope into which project minute echinulations, olivaceous in KOH, rusty brown in iodine, witl an eccentric oil drop, apedicellate or with only a broken stump of a pedicel; capillitium of frequently branched threads, tapered to their apices, many rather crooked or with sinuous walls, olivaceous brown in KOH, rusty brown in iodine, thick-walled, main branches pitted. Habit, habitat, and distribu tion.-Gregarious in a spruce plantation in the Saginaw Forest, Ann Arbor, Waslitenaw County, October 16, 1936 (Smith-6o9o). The description is drawn from this collection. Amnong Kauffman's collections are 2 which belong here, 1 from Houghton, Houghton County, which is immature, and 1 from New Richmond, Allegan County. It apparently fruits during the late summer and fall, and appears to be rare. Discussion.-Kauffman considered this form to be the same as L. polymorphum, but Lloyd's statement that it is the true L. polymorphum of Europe is open to question, since my collection is typical of L. polymorphum f. cepaeforme in shape. My collections of f. cepaeforme were white when young. This is a field character of some significance in this case, because of the extremely brilliant color of L. coloratum. Consequently, I

Page  52 52 PUFFBALLS AND ALLIES IN lICHIGAN believe that it is desirable to maintain tle concept of Coker and Couch for this species. It may be objected that yellow forms of the L. umbrilnum series are lumped with white forms in the varieties recognized in this paper. But further studies need to be made on the umbrinum group, in so far as the yellow fruiting bodies are concerned, to determine how much of the yellow coloration reported is caused by the color changes accompanying tlhe maturation of the specimens and how much is caused by the actual presence of yellow pigment in the inner wall and the scurfy covering of immature specimens. My notes are incomplete in tlis respect, and I have found notling in tlle literature bearing upon it. LycopIerdon p)usillum Persoon (P'1. X, Fig. 2) Fructification globose to subglobose, 0.8-2(2.3) cm. in diameter, pinclied off at base and witli a short rooting base or only a few brancled wllite rhizomorphs projcctilng, white whmen young, surface of spore case obscurely flocculose, the flocculent layer in some becoming aggregated into a dense covering of very small warts or granules wlhiclh separate from each other in drying or by breaking up into minute areolate patches wlhich finally sllrink to flakelike particles before falling off; surface pale brown to dark dull brown at maturity and obscurely spotted with paler spots in many specimens; gleba passing through yellow to greenisll yellow and finally olive brown to deep coffee brown, occulpying tlhe entire fruiting body; sterile base none; apical pore slow to form but finally rather large (3-5 mm.) and with lobed margins. Spores globose, 3.5-4.5 p in diameter, witli a thin hyaline envelope into which mniute eclinulate processes project, apedicellate or witll a broken stump of a pedicel, with a large usually eccentric oil drop; capillitiuml of tlick-walled, yellow-brown threads wlhicli are often ratler crooked or witl sinuous walls and tapered gradually to the apex, tapered portions paler than tlhe tlicker parts, 2.5-4 ((6) in diameter, the largest threads witl pitted walls.

Page  53 LYCOPERDALES 53 Habit, habitat, and distribulion.-Densely gregarious to subcespitose, or occasionally scattered, on thin soil, along roadsides, on waste land, pastures, and the like. It fruits from June to late fall depending on the weather. It occurs throughout the state. My collections are all from the vicinity of Milford, Oakland County, and Waterloo, Jackson County. Povah collected it on Isle Royale, Keweenaw County, and Potter found it abundant in the vicinity of Ithaca, Gratiot County. Discussion.-Coker and Couch commented on the fact that the scurfy outer coating does not slough off as a superficial layer. This is exactly as I find it in my specimens, though the particles finally disappear, leaving the paler spots mentioned in the description. Most mature specimens which appear to have lost the outer covering will, upon examination under a lens, be seen to have the remains of the flakes and granules dried down to very minute adhering particles of the same color as the surface. The olivaceous gleba, small size, peculiar surface covering, and lack of any sterile base are the important characters to remember. The spores and capillitium are not diagnostically important because they resemble those of numerous other species. Lycoperdon oblongisporum Berkeley and Curtis (1'1. X, Fig. 3) Fructification 1-3 cm. in diameter, globose to flattened, attached by a persistent rhizomorph, white when immature, in age darkening to near "warm sepia" (Ridgway); surface covered by a thin layer of fibrous-granular material (the outer peridium) which breaks up in an irregular manner into minute patches that in turn become grouped to make up an obscurely areolate pattern and eventually slough off, at least around the mouth, to expose the smooth more or less wood-brown endoperidium; mouth nearly round and usually with a lobed to crenate margin, rather large; gleba yellow to olive brown and finally very dark olive brown; sterile base greatly reduced and chambers not conspicuous. Spores 5-6 x 3.5-4 p in diameter, narrowly to broadly ellip

Page  54 54 PUFFBALLS AND ALLIES IN MICHIGAN soid, apedicellate and olive-yellowish under the microscope in KOH, smooth but with a thin hyaline gelatinous envelope, very obscurely punctate as seen under an oil-immnersion lens; capillitium of flexuous to very crooked threads witl thickened walls, 3-5 p in diameter, branching infrequent, cross walls very rare, olive yellow, near Isabella color (Ridgway) as mounted in KOH, tapered evenly to long-drawn-out apices. Habit, habitat, and distribution.-Gregarious on exposed soil, often around stumps or bases of trees, July 5 to August 1 2 in the vicinity of Ithaca, Gratiot County. Potter made over 5 collections during the season of 1948. His specimens are the only Michigan collections which I have examined. Discussion.-Coker and Couch commented on finding collections with spore characters intermediate between this species and L. pusilllum. Lycopcrdon pusill2um is an extremely common fungus in Potter's collecting grounds and it has been possible to make a careful comparison. The 2 species do resemble each other very closely in all except spore characters. In nearly all collections of L. pusillum the spores are globose and distinctly echinulate, whereas in L. oblongisporun the spores are truly ellipsoid and no markings can be seen on the spores except with the very highest power optical system. L. oblongisporum was originally described from Cuba and has been considered rare in the United States. Its discovery in Michigan is of great interest. Lycoperdon polymorphum Vittadini f. cepaeforne Lloyd Fructification globose, pear-shaped, or with a short stemlike base and enlarged above into a globose to depressed-globose head; base rooting or merely with several rhizomorphs; 1-3.5 cm. broad in widest point, up to 4.5 cm. tall, white when young; surface of spore case covered by a thin coating of very short spines, which in age may become separated into minute warts or granules before disappearing; granulose layer (exoperidium) in age similar to that of L. umbrinun vtar. umbrinumn and breaking up in the same way; surface olive brown at maturity and glabrous only in patches; apical pore round or nearly so;

Page  55 LYCOPERDALES 55 gleba changing from white through yellowish olive to dark olive brown, occupying most of the fructification; sterile base only slightly developed and lacking distinct chambers. Spores 3.7-4.2 p in diameter, globose, olive brown in KOH, rusty brown in iodine, with a thin hyaline envelope into which minute echinulations project, usually with a broken stump of a pedicel; capillitium much branched, main branches 5-7 P in diameter, others 3-4 p, tapered to apices, olive brown in KOH and rusty brown in iodine, walls thick and pitted in large branches. Habit, habitat, and distributionl.-Densely gregarious to scattered. It fruits in the fall in the vicinity of Ann Arbor, Washtenaw County. Johnson reported it in July and August in Ohio, and stated that it was frequent. I have seen it on humus in the woods and on sandy soil in waste places. Discussion.-The larger size and more granular nature of the superficial coating distinguish this species from L. pusillum. The sterile base in the Michigan collections is poorly developed. The typical form is said to have a well-developed sterile base in which the cavities are visible only with the aid of a lens. Typical specimens of L. pusillum and L. polymorphum can be distinguished from each other by their difference in size, and by the presence of a sterile base in polymorphtum and a flocculent rather than warty or spiny surface covering in p/usillum. Lycoperdon marginatumn Vittadini (PI. XI, Figs. 1-2) Fructification 1-4(5) cm. broad, globose to subglobose at first but becoming somewhat flattened and when mature nearly always broader than tall, usually plicate or sulcate on underside and narrowed to point of attachment; some with a short rooting base; surface of spore case at first covered with a thick coating which becomes blocked out into warts with sharp tips or into compound warts with the tips cohering, the whole layer white and remaining so or finally discoloring slightly, at maturity flaking off in patches or sleets to expose the smooth to obscurely pitted, pale to dark olive-brown surface, the warts more poorly delimited toward margin and on under side of fruiting body,

Page  56 56 PUFFBALLS AND ALLIES IN MICHIGAN separation of warty layer usually beginning along a median line between tlhe stem and top, tle part covering the under or lower side usually persistent; surface finely furfuraceous to unpolished immediately after cortex lIas sloughed off but finally becoming polislled, furfuraceous particles yellowish to dull brown; gleba olive to grayisl brown but pallid after spores have been shaken out; sterile base well developed, chambered, clhambners relatively large (i mmnn.~), in some arched in tlhe center and extending up the sides as a thin layer, in others poorly developed and occupying only tlhe base. Spores 3.5-4.2 p in diameter, globose, smooth (under an oilimmersion lens appearing minutely punctate from0 pores throtugho tle wall), witl a very thin lhyaline surrounding sheath and a broken stub of a pedicel, pallid olivaceous to brownish in KOH, rusty brown in iodine, with a distinct oil drop; capillitium of slender straiglht to contorted fi!aments 3-6 p in diameter with thickened pale to dark yellowish brown walls, in iodine dark rusty brown, paler and tapered toward tlle apex. al)it, halbital, and distribultion?.-Cespitose-gregarious or at times solitary in sandy waste ground, pastures, fire lanes, and open oak woods on exposed soil; from July on to the end of the mushroom season but usually most abundant in September. It is a common Mlicligan species and occurs throughout the state. l)isclssio7.-Tlis species of Lycolerdon is very easy to identify in tlle field, if mature specimens are at hand, because of tlle manner in iwhiclh the tlhick layer of warts (the cortex) separates from thle surface (PI. XI, Fig. i). As Coker and Couch pointed out, L. Curtisii and L. marginatum are difficult to distinguish 1whlen immature, because at that stage the cortex in both looks very ll:.hl alike. In L. CTurtisii, however, tlie cortex does not sloughl oi, so mature specimens can be separated by this character and also by the different type of capillitium. Lycopcrdlo pyriformnc Persoon (P1. XII, Figs. 1-2) Fructification pyriform to slbgloblose,. 5-3(4.5) cm. in diameter, broad in widest part, 2-3(4.5) cm. high, typically not

Page  57 LYCOPERDALES ) / wrinkled or plicate at juncture of the enlarged part with the stipelike base; surface of spore case pallid to tawny brown in immature stage, typically darker rusty lrown at maturity but in some the smooth surface (beneath the granules) yellowish; outer coating at first smooth at least ovecr the rounded part but soon breaking up into areolate patches, which redivide into smaller units and in drying form minute granutles or particles; at times cortex at least over apex composed of small spines and granules, remains of cortex relatively persistent and rough to the touch like a file, eventually falling away over fairly large areas exposing the smooth endoperidium; apical pore slow to form but in age sometimes forming an irregular slit 1 cm. or more long; gleba white wlhen ilmmature, becoming olivaceous and finally deep olive brown, occasionally with a grayish cast in overmature specimens; sterile base often only sliglhtly developed (in globose forms) or occupying tle entire stemlike base (in stipitate forms), chambers small; base connected to substratum b)y numerous white rhizomorphs. Spores globose, 2.8-3.5 p in diameter, smooth, with a thin continuous hyaline envelope (as seen under an oil-immersion lens), apediceliate, with a large oil drop, pale olivaceous to deep olive brown depending on degree of maturity; capillitiumi of sparingly branched threads, even to flexuous, the cells tlickened and olivaceous to dull brown (in KOH), bright rusty bro-wn in iodine, tapered to the tips and paler in color in tapered portion, 3-6 p in diameter. Habit, habitat, and listribution.-Ccspitose to scattered, or occasionally single. It usually occurs on or around rotten hardwood logs and stumps, around old sawdust piles, or on humuls very rich in ligneous debris. It is a very common 'uingtls throughout the state. The old fructifications can be found any timle during the year, but the fruiting period typically is froJm early September to the heavy frosts of late fall. Old weathered spore cases may persist through tlhe winter and on into the following summer. Discussioz.-This species is considered a fairly good esculent and has the added advantage of occurring in sufficient quantit)y

Page  58 58 PUFFBALLS AND ALLIES IN MICHIGAN as a rule, to allow one to get Celough lor a lecal. As it is tlhe only common Lycoper(don in Mlicligan on decaying wood, the habitat is a fairly reliable field character. The scales may or may not be present over tlle rounded part of immature specimens (those which one would be seeking if collecting for the table), as may be seen from the illustrations. Lycopcrdon m nscormll Mlorgan Fructification pyriform to g(lobose or depressed-globose but contracted below into a slort stemlike base, 1-2 cm. broad and lip to 2.5 cm. higl, fine white rhizomorphs extending out from base; surface of spore case at first covered (especially over apex) with short broad cones and distinctly smaller granules (larger (coneS somewhat resembling those of L. geimmatuln but nmuch smaller), narrowed part mierely covered by fine granules; granulose-spiny coating very persistent but finally wearing away leav'ing a smlootl yellowish to olive-brown surface; gleba olive bro)-wn; sterile base greatly reduced in globose forms, occupying stalklike part whenl the fructification is pyriform, chambered but tlie chambers small. Spores 3.5-4.2 p, globose, pale olive brown in KOH, rusty brown in iodine, pedicels broken off and littering the mount, surface witlh a very thin hyaline envelope into which project slort ecllinulations; capillitium of threads 3-5 p in diameter, tlhe walls olive brown in KOH, fulvous in iodine, somewhlat tickened and somiewhlat pitted, occasionally septate, threads cylindric to sinllous or some with irregular thickenings, witll a tendency to break iup if pressure is applied to cover glass of mount. Habit, habitat, and distribution.-A very rare species as far as my experience goes. I have 1 collection from Mud Lake Bog, Waslitenaw County, Septe 1, 945 (Smith-20589) whicl appears to belong ]lere. The description is taken from it. DiscussliTio.-The specimens I collected grew on sphagnum withl somec Polyltichulmn scattered about, but it was plain that tlIe fungus llad no particular affinity for the Polytrichum. To judge from the accounts of the fungus in the literature the

Page  59 LYCOPERDALES 59 same great variation in length of stipe occurs in this species as is frequently encountered among agarics growing in a similar habitat. Consequently, I am inclined to place no taxonomic significance on the greatly elongated sterile base of some specimens even though it is a very striking field character. During some seasons this fungus is very likely to be abundant in the northern part of the state. Its qualities as an esculent are very poorly known. Lyco/perdon Peckii Morgan "Plants shaped as in L. gemnmaturn, the distinct stalk up to 3 cm. long; the mycelium ropy; peridium about 1.5-3.8 cm. broad, covered with tapering spines about 1-1.5 min. long, which as the plant grows split at base into 2-4 parts which remain united at their tips and are easily rubbed off, leaving pale, smooth, circular spots which are surrounded by minute granular warts and dots; color of the longer spines in youth and until near maturity a delicate and pretty purplish lavender, at maturity changing to buffy brown. Toward the base the spines become more slender and hairlike and are more obviously intermingled with granular matter. Except for the lavender color of the upper half, the plant until near maturity is nearly white. Subgleba composed of rather small cells and occupying about a third of the plant. "Spores (of No. 5749) olivaceous brown, spherical, 3.6-4.4 p thick, with a small oil drop and a short pedicel, the wall obscurely dotted (pitted or minutely warted) and appearing faintly striate radially in optical section; surrounding the spore is the thin hyaline layer that is found in most Lycoperdon species. Capillitium threads very slightly branched, up to 6 p thick, and tapering to delicate tips" (Coker and Couch, 1928: 84). Discussion.-A single collection (Smith-21538) from the University of Michigan Biological Station at Douglas Lake, Cheboygan County, probably belongs here. The spores are globose, 4-5 p in diameter, usually with a short broken pedicel, and have a very thin hyaline envelope into which short rodlike processes project. The capillitium is not distinctive, being composed of

Page  60 6o PUFFBALLS AND ALLIES IN MICHIGAN thick-walled brown threads sparingly branched and J3-6 P in diameter. The spines (remains of the exoperidium) were brownish when the fructification was found. The covering of spines and granules is to some extent like that of L. perlatum, but one would hardly confuse the 2 species in the field. As Coker and Couch suggest, however, they are closely related. A study of more material is highly desirable. Lycolperdon j)erla un m Persoon (Pls. XIII, XIV, and P1. XV, Fig. I) Fructification variously shaped but typically obovoid to turbinate and possessing a stemlike base, (1.5)2.5-6(() cm. across the widest part, narrowed abruptly below this part and often plicate to merely wrinkled along the edge and downward for unequal distances on the stemlike base, sometimes merely pear-shaped, base 1-2.5 cm. broad where attached to substratum; typically white when young and fresh, but spines not infrequently brownish in age; surface of spore case covered above:by many short cones (spines) with round bases and pointed apices, between these occur many more shorter warts or very small cones, or the surface between larger cones may be furfuraceous, the larger cones soon deciduous, leaving pale round spots on the surface (particularly around the pore), in age the surface at times becoming almost smooth from the falling off of both spines and granules; coating of spines and granules extending down onto the stalk but difference in size between them becoming less apparent or at least the larger spines more widely distributed, the smooth surface (in age) yellowish to dark avellaneous; gleba white when young, changing through yellowish above to olive brown (in the literature occasionally reported as tinged purplish in very old specimens); sterile base occupying the stalklike region, witli clhambers up to 1 mmn. or sliglhtly more in diameter, white throughout at first, but in age often olive-brownish to chocolate-colored; pseudocolumella elliptic. Spores 3.5-4.2(4.5) p in diameter, globose, apedicellate, pale to dark olivaceous brown in KOII (depending on degree of maturity), yellow to deep rusty brown in iodine solution, minutely

Page  61 LYCOPERDALES 61 ecllinulate, the c(liiinllations projecting to the surface of a distinct thin liyalinc covering, capillitium of thick-walled, simple to sparsely l)ranched filaments yellowish to dark brown (depending on degree of maturity), walls smooth and not or only very slightly pitted, filaments flexuous to crooked, 3-7 p in diameter, tapering to 1.5-2 p at free ends. Habit, habitat, and distribution.-Single, gregarious, or cespitose, often in arcs or "fairy rings" on humus in conifer and hardwood forests tllroughout the state. Its typical fruiting period is late Alugust to late October, depending on the season. The most luxuriant fruiting I have seen was early in October in a local conifer plantation. Povah found it on Isle Royale, Keweenaw County, July 29, 1930. Discussior7.-This and L. pyriJorne are the commonest puffballs in the state. The spotlike scars marking the position of the large deciduous cones are the most reliable field character of mature fructifications, and the round pointed cones themselves serve to distinguish immature specimens. Here in Michigan the typical white form of thle species is the common one, though in the north a form with dark wood-brown spines and granules should be seen occasionally. L. perlatuln is Imuch more widely known under the name L. gemm.atnuwn Batsch. Lycoperdon pediccllatum Peck (1P1. x, Fig. 2) Fructification variable in shape, 2-4 (5) cm. broad at widest part, 3-6 cm. tall, with a thick stipe terminating in a flattened head, pyriform or globose to subglobose, attached to substratum by numerous thin white rhizomorphs; surface of spore case covered over upper part by spines 1-2 min. long, single, or in groups of 3 to 4, then with converging tips, at first crowded and forming a continutous covering but slhrinking on drying to expose a smooth polished surface in mature specimens, gradually falling away to expose tlhe obscurely pitted to reticulated surface, which is dull brown to grayish; spines smaller and more granulose material is present toward tile broad rim of the fructification and downward to the point of attachment, sometimes pli

Page  62 62 PUFFBALLS AND ALLIES IN MICHIGAN cate around the base; gleba grayish olive to dull cinnamon brown depending on degree of maturity; apical pore most often slitlike or an irregular tear; sterile base well developed and occupying the stipelike base in elongated specimens, considerably reduced in globose or subglobose specimens, with chambers up to 1 mm. in diameter, but usually smaller, pallid to dull brownish. Spores subglobose to broadly oval, 3.5-4.5 x 4-5 p in diameter, pale cinnamon brown in KOH, dark rusty brown in iodine, with a very thin hyaline envelope and fine echinulations projecting into it, pedicels persistent, hyaline, with thickened walls, often flexuous, io-i8 p long, capillitium of brown, thick-walled, sparingly branched filaments 4-6 p in diameter. Habit, habitat, and distribution.-Scattercd to gregarious on very rotten hardwood logs and stumps or humus rich in ligneous debris. It usually fruits during September and October and appears to be fairly abundant in seasons of ample rainfall. E. B. Mains collected it at Harbor Springs, Emmet County, in 1931 and both Mains and I have seen it at the Edwin S. George Reserve near Pinckney, Livingston County, and in neighboring wooded areas. Kauffman reported it throughout the state, but the only collection of his which was so labeled, turned out to be Lycoperdon umbrinum. Discussion.-In the Michigan collections there is a tendency for the membranous layer (endoperidium) to be rather brittle and break up as is characteristic in Calvatia, but the tendency is not pronounced enough to justify removing the species to Calvatia. It was formerly placed in Bovistella by some authors, but I am in agreement with those who retain it in Lycoperdon. Lycope rdon un mbrinunm Persoon var. umbrinum (P11. XVI, Fig. i) Fructification (1.5)2.5-5 (9) cm. broad at widest point, 3-8(12) cm. high, typically eve nly enlarged upward or pear-shaped, occasionally nearly globose, often sulcate to plicate beneath the enlarged upper part; base 1-4 cm. broad, white to yellowish or at times lemon yellow wihen immature; surface of spore case

Page  63 LYCOPERDALES 63 typically covered with a thin scurfy or granulose coating mingled with very slender short spines which are erect or (over stemlike base) flattened toward each other, this coating relatively persistent or finally breaking up into concentric zones (on base) or areolate patches (over top) and eventually disappearing leaving the smooth yellowish to dull-brown surface exposed; gleba white, when immature, changing through yellowish to brown, grayish brown, or purplish brown in age, not infrequently retaining an olivaceous cast; pseudocolumella absent or only slightly developed; sterile base occupying the entire tapered part of fruiting body and extending as a thin layer up to the widest part, chambered, chambers i mm. ~ in diameter. Spores 3.7-4.7 p in diameter, globose, pale chocolate brown revived in KOH, deep rusty brown in iodine, verrucose; warts truncate, crowded and of hyaline material (yellowish in iodine), in KOH the warts fusing somewhat but not losing their identity (Coker and Couch reported the spore as appearing to have a thick hyaline wall after being soaked in water); capillitium of threads 3-5(8) p thick, branched, walls dark brown, thickened (2 up ), pitted, even or flexuous to crooked or subcontorted, tapered to apices, often with irregular enlargements and constrictions. Habit, habitat, and distribution.-Solitary, gregarious to subcespitose, on leaf mold and needle beds under conifers. September to late October is its typical fruiting period, and it is to be expected throughout the state. It is very abundant at times in the Saginaw Forest near Ann Arbor, Washtenaw County. Discussion.-One collection (Smith-4936) represents the form which is lemon to golden yellow when young; another collection (Smith-5o63) has the form (here considered typical) which is white to pallid brownish when immature. The variations within this species have presented a problem to all who have studied it. Lloyd's treatment of it indicates that he, in spite of his strong statements in regard to variation in it, failed to arrive at a satisfactory species concept. Lloyd's illustration (3: 438, Fig. 250) of variation in the cortex completely fails to impress anyone who has seen a fairy ring of a Cortinarius superimposed on one

Page  64 64 PUFFBALLS AND ALLIES IN MICHIGAN of a Clavaria to such an extent that the fructifications alternated throughout most of the ring. Lloyd believed that because 2 fructifications of Lycoperdon grew close together, even though they differed sharply in the character of the cortex, they came from the same mycelium and hence "were the same plant." Had he illustrated a series of intergrading forms, his opinion would have more weight. He apparently failed to realize that le, as well as the "learned professors," worked largely on circumstantial evidence and that a chance occurrence near together of 2 closely related, but possibly genetically distinct, taxonomic units is by no means sufficient evidence for assuming that they are identical. The treatment by Hollos (1904) is the one I have followed here in the main, for my collections seem to fall readily into the units he established. The following varieties in addition to var. umbril nu are recognized. L. um brinum var. atropuirpurcum (Vitt.) Hollos (I1. XVI, Fig. 2) Fructification 2-5 cm. broad at widest point, depressed-globose or subpyriform, often sulcate at base and furnished with a mycelial rootlike process (particularly when growing in sandy humus), 2.5-6 cm. high; surface covered with a dense coating of granules and hairlike spicules which become worn away in age or only remain over the narrowed lower part; sterile base usually occupying lower one-half to two-thirds of fructification; olela soon distinctly purplish brown. Otherwise as in var. unlbri,um. Habit, habitat, and distribut ion.-Typically solitary in sandy oak woods in the southern part of the state. It is usually found late in September or October, and is common in very wet seasons. L. lumnbrinnum var. floccosum Lloyd (Pls. XVII-XVIII) Fructification globose to pear-shaped or with an elongated stalk, (2)3-8(o1) cm. broad at widest part, (3)4-10(12) cm. high; surface covered to a varying extent with long flocculent spines,

Page  65 LYCOPERDALES 65 in addition to spines like those described for var. umbrinum; sterile base, spores, and capillitium as in var. umbrinum. Habit, habitat, and distribution.-Solitary, gregarious, or cespitose, often in fairy rings in October; abundant in a local spruce plantation in the Saginaw Forest, near Ann Arbor, Washtenaw County. Discussion.-As in var. umbrinum a yellow form occurs, but in this variety the typically white form and the yellow have numerous intergradations, and the white form is likely to turn yellowish before maturity if badly rain-soaked. There are still other variants of L. umbrinum in the area which need to be studied. One is close to var. astcrospermum, but is lilac over the upper surface when immature. Lycoperdonl rim ulatllm Peck (1'1. XIX, Fig. 1) Fructification 1-3(5) cm. broad, globose to depressed-globose and usually deeply plicate around the pointed fibrillose rooting base; surface at first whitisll to grayish or finally pale drab, smooth and unpolished (like kid) at first but soon indistinctly rimose-areolate, becoming more distinctly rimose-areolate in maturing and finally the areolae become compound, that is, the larger blocks are further divided into more indistinct smaller units and tlese in turn are obscurely dotted, the arcolate layer finally separating from the surface of the spore sac as thin plates or scales; surface of spore sac smooth, purplish gray to brownish, opening by an apical pore; gleba whlite when immature, changing through light yellow to gray and finally to deep brownish purple (benzo brown of Ridgway); sterile base occupying the rooting base and extending along tlhe under side of the gleba to about the broadest dimension of the fructification, chambers relatively large and distinct. Spores globose and pedicellate, 5.5-8 p in diameter, covered with colored warts embedded in a thin hyaline envelope, with a large oil drop, dark rusty brown in iodine, pedicel 2-3 (rarely more) times as long as the diameter of the spore if not broken off, its walls thickened, often minutely irregular to wavy, and in iodine pale yellowish; capillitium of brown-walled threads

Page  66 66 PUFFBALLS AND ALLIES IN MICHIGAN (rusty brown in iodine) with thickened walls which are almost nodulose in places, tapering gradually to apices, 3-7 p in diameter, sparingly septate. Habit, habitat, and distribuItioni.-Closely gregarious in a sandy meadow under a tree, Ithaca, Gratiot County, collected by Victor Potter, September 13, 1947. Potter made a collection of some immature specimens in tle same area in July of that year, which are apparently this species, and also made additional collections in 1948. Lloyd reported it from hMiclig-an, but I lhave not seen his specimens. There were none among Kauffman's collections. Discussionl.-Potter's collections of mature specimens lhad no sign of tle reddish brown mentioned by Coker and Couch, but the puffballs may not have been old enough. The species is generally reported as rare. It is one of the easiest of the genus to recognize in the field, if the dotted areolae representing the remnants of the outer covering are still present. In the character of its spores and capillitiun it is perfectly typical of Lycoperdotn. Lycoperdoii pulcherri7mum Berkeley and Curtis (P1. XIX, i:ig. 2) Fructification body 2-5 cm. -broad in widest part, 3-4.5:m. highl, pear-shaped to subglobose above a narrowed often plicate lase, attached by white rhizomorphs; surface of spore case at first covered by a dense coating of very long (4-6 mm.) slender white spines (they remain whitish even when dried), which become arranged in fascicles or cones by being united at the tips, spines finally falling to expose the smooth, typically dark purplebrown surface; pore with a torn to lobed margin and becoming large; gleba white, clhanging tlrough yellow to dark purplebrown; sterile base occupying tlhe basal one-third to one-half of the fruiting body, clambers distinct, but typically less than mi m. in diameter, purple-brown in age. Spores globose, 4-4-5 p in diameter, with a lhyaline surrounding layer into which short spines project, broken pedicels littering the mount buIt spores retailing a pedicel 1o-13 p long; capillitium lbranched, sliglitly pittcd, tlickest cells 6-7 p in di

Page  67 LYCOPERDALES 67 ameter, dark brown, tapered to slender flexuous or contorted almost hyaline apices. Habit, habitat, and distribution.-In spite of reports in the literature to the contrary, I doubt that L. pulcherrimum occurs in Miclligan. Coker and Couch (1928: 73), who described and illustrated it exceptionally well, stated that "we never find it in abundance, and often only a single specimen at a time." It is to be sought for in the hardwood forests and brushy areas of the soutlern counties. Discussion.-Kauffman's 5Michigan collections identified as this species proved to be L. echinatum or L. umbrinum var. floccosuni. My description and illustration are taken from material collected near Cisco, Texas, by E. A. Smith in September, 1935, and on January 6, 1937. I am not certain that the January date is significant, for the specimens were mature and may have been standing for some time. All of my collections from \Iichigan are referable to L. echinatum. I have not seen Lloyd's records from the Great Lakes region, but since he reported it as one of tile characteristic species of the region I have included it here. J. A. Stevenson informed me that there were no \Iichigan specimens in the Lloyd Herbarium. It must be kept in mind that the species was originally described from the soutlheastern states and that there is a substantial difference between the species wlhich inhabit tlat region and this. I have examined what I consider to be typical specimens from Kansas as well as from Texas. Lycopcrdon cchilnat unt Persoon (P1. XX) Fructification 2-4(5) cm. broad in widest part (excluding spines), globose to subpyriforin when immature, becoming somewhat flattened, pinched off to a base 5-8 mm. broad and with white rlizomorphs penetrating the substratum; surface covered with a dense coating of spines 3-6 mm. long and soon arranged in groups withl more or less convergent tips; spines white when young, in age becoming dark brown and falling off, leaving the surface reticulated, reticulum formed of minute

Page  68 68 PUFFBALLS AND ALLIES IN MICHIGAN scurfy particles and eventually wearing away, leaving the shining pale to dark purple-brown surface naked (surface of incompletely ripened specimens often pale brownish); gleba distinctly purplish when fully mature, white when young, then yellowisll to olivaceous, changing further to grayish brown and finally purplish, in some with a yellow zone persisting above the sterile base; sterile base not well developed, the chambers small and in some indistinct, extending up the sides for some distance as a narrow zone, purplish gray to brownish. Spores globose, 4-4.5 p (Smith-20563), 4.5-6 p (Smith-20731), 4.8-6 p in diameter according to Coker and Couch, with a surrounding envelope of hyaline material into which short warts project, long hyaline pedicels usually broken off and littering the mount; capillitium of tlick-walled dark brown filaments tapered to the apices, tapered part paler to hyaline, threads branchled, pitted, often crooked to almost contorted or merely witll irregular swellings and constrictions, tlickest threads 6-8 p in diameter. Habit, habitat, and distribution.-Single to gregarious on humus or very rotten wood in hardwood forests. It fruits in late September and October during seasons of ample rainfall and is often abundant locally. Discussioin.-Tlhe immature stages of this species are a rather beautiful siglit in the woods, but as specimens age and the spines darken the fructifications become relatively inconspicuous against a brown background of dead leaves and dead wood. Immature specimens of L. echinat1um and L. pulcherrimum are difficult if not impossible to distinguish from each other, but tllis will cause no inconvenience to tllose collecting for the table, since botli are edible. In the mature stage the surface of L. cchinatumn is reticulate, but that of L. pulcherrimum is not (compare P1. XIX, Fig. 2 and P1. XX, Fig. 2). For further comments, see L. pulcherrimum. Disciseda Czerniaiev Fructification obovate to depressed globose, peridium 2-layered; outer layer (exoperidium) membranous or a sand case,

Page  69 LYCOPERDALES 69 fragile, deciduous, usually remaining as basal disk; inner layer (endoperidium) papery but tough, opening either by a basal or apical pore (stoma) depending on the species; sterile base none; capillitium present; spores colored, roughened. There are 3 species in Michigan, distinguished primarily on 3 ranges of spore size, and all are apparently very rare. KEY TO SP'ICIES OF Disciseda i. Spores 3-5 /u in diameter, smooth or nearly so....................D. candida i. Spores 6-8/ in diameter, distinctly warty...................D. subterranea i. Spores 10-12.5 g in diameter.................................D. Muelleri Disciseda candida (Schw.) Lloyd ('1. XXI, Fig. i) "Plants single or cespitose, compressed globose, with a single rhizomorph-like root when young and fresh; 0.5-2 cm. tlick, 2-3.5 cm. wide before maturation; o.8-1.5 x 1.5-2.5 cm. when dry; growing in tile soil and partly exposed when fully grown; surrounded until maturity by a thickish cortex (outer peridium) the upper and thicker part of which remains attached to the inner peridium by a layer of spongy fibrous material, the lower part tearing irregularly from the upper and separating more or less completely from the inner peridium and remaining in the ground as broken and inconspicuous fragments. The upper part with the inner peridium attached is now free and is easily knocked out of the cup-like lower part by rain, etc., turning over so that the exposed part of the inner peridium is now above. A small torn hole now appears at the place of attachment of the stalk, in the center of this upturned base, and the spores begin to escape. The cortex is a sand case held together by a woven white mycelium. The inner peridium is rather firm and rigid, pale brown then silvery gray to slate gray in age, minutely granular or scurfy on the exposed (lower) part, densely scurfy above under the cortex. This thick spongy tissue above and thin granular layer below are the remains of a watery translucent layer, o.6-i mm. thick which lay between the more fibrous layers in youth. The inner peridium is very indistinct in sections when fresh, becoming conspicuous on maturing and drying. The whole

Page  70 70 PUFFBALLS AND ALLIES IN MICHIGAN plant shrinks to scarcely more tlan half its original size upon drying. Glebal chambers extremely minute, 5-25 J wide, but often longer; no definite tramal plates are formed, as in Lycoperdon and Sclcroderma, but tlle basidia arise from irregularly anastomosing strings of tlreads and the chambers are very irregular. Gleba white when fresl, changing as the plant matures through yellowish olive to brown, at times faintly purplish. Thlere is no sterile base. "Spores (of No. 5967) brown, at times with a tint of purple, globose, 3-6.4 p1 thick, warted, and witll a short pedicel (mucro). Capillitium threads 3-5 p thick, irregular, not rarely branched, extending inwards from the walls of the peridium; after maturity breaking up into short pieces. Basidia (of No. 6092) shortpyriform, 6-7.4x9-14 P, usually with 4 slender apical sterigmata of equal length" (Coker and Couch, 1928: 1 )9-140). Discussioni.-Thc only material I have seen from this state, aside from abundant collections by Potter from near Ithaca, Gratiot County, consisted of a single fruiting body from a sandy field near Steere's Swamp, near Ann Arbor, Washtenaw County, collected by A. II. W. Povah, October 27, 1914. Perlhaps tile best field character for this fungus is its likeness to an acorn. The resemblance in sliape is striking in the mature specimens. Tlie important feature to keep in mind, however, is tlat tlhe pore is at the base, not thle apex, of tlhe spore sac. The spores in tlhe Micligan collections are nearly smooth. For furtlher comments, see D. subl)crralca. Disciscda sublcrran-caa (Pk.) Coker and Couch ('1. XXI, l'ig. 2) Fructification 1-2 cmn. broad, depressed globose to distinctly convex-flattencd, at maturity appearing acorn-like because of thle way the spore sac rests in tle remnains of tlhe outer layer which is a mixture of sand and llyplae, usually opening by a round pore with elevated, entire to somewhlat lobed margin; surface of spore sac unpolished, with a bluisl to glaucotus gray coating over a dull brown ground color; gleba dull rusty cinnamon. Spores globose, 6-8 p[ in diameter, coarsely echinulate with hyaline projections, apedicellate; capillitiuml pale but with a

Page  71 L YCOPERDALES i71 brown tinge, flexuous, readily breaking up, walls slightly thickened, cross walls and branches rather numerous. Habit, habitat, and distributlionl.-Gregarious in sandy pastures, summer and fall; Riverdale and Sumner, both in Gratiot County. All collections were made by Victor Potter. Discussion. —Potter obtained all 3 species of Disciscda reported lhere. Disciscda sll)tcrralna and D. candida, to judge from Potter's collections, look exactly alike. It has not been verified as yet whethcr the manner of development described by Coker anld Couch for D. canllida applies to this species or not. Disciscda Mlccllc'ri (Berk.) Cunningham Peridiutln up to 2.5 cm. diameter, subglobose, witl a short stout rooting base; exoperidilum soon umber, with minuite pale subpersistent warts; cndoperiditum firm and rather thick, brownl. Gleba reddisll brown, capillitium tlreads flaccid, pale, lbut little branched; spores globose, 10-12 pj diameter, epispore rcddisli umber, coarsely spinulose, apcdicellate. Habit, habitat, and disltribtltion.-Potter found an old fructification, which appears to belong lhere, in a sandy pastture near Riverdale, Gratiot County, on ()ct. 13, 1948. Discuission. —The microscopic cliaracte-rs of Potter's collectioil are as follows: Spores globose, 10-12.5 u, with coarse closely set hyaline warts, dark brown beneath the warts and wall distinctly thickened, usually fturnishcd witl a short stum!np o! a pcdicel, capiilitium1 of flextotls very pale tlreads whllichl blrak tip readily into relatively short pieces; walls slighltly tllicklclcd, branlcliing only occasionally, cross walls rare. No rl1ixoi(1o(pll OF0 rooting base was present on the Potter collection, but a basal scar indicates that one was very likely present originally. lThec pale flexuous capillitiuml and large spores of his specimen also clheck for tllis species. Co)nse(lcnltly, I have reported it here under tllis name, althoughl I do not regard tle identification as necessarily final. Whien one considers that such a distinctive 1ungus as Titlo.stoma slriallnl lias just as peculiar a distribution, it is not justifiable to rule D. MI/llcri out of consideration oi tlat basis. Both wvere described from Australia.

Page  72 72 PUFFBALLS AND ALLIES IN MICHIGAN Bovistella Morgan Fructification subglobose to turbinate, remaining attached to substratum; surface covered by a dense floccose somewhat persistent coating which usually becomes aggregated into slender spines or particles but may be merely granulose; wall of spore case delicate, opening by an apical pore and collapsing as spores escape; sterile base present, persistent; capillitium of free, mucl branched units with acute ultimate branches, the main brancles tliicker than the diameter of the spores; spores globose to oval, smootlh or echinulate, with a thin hyaline envelope. Cunninghlam considered Bovistella to be a synonym of Bovista. I lhave followed American investigators in recognizing it, but do so witl the distinct impression that at best it is a very weak genus. KEY TO SI'I:CII'S OF Bovistclla i. Fructification small (5-8 mlm. in diameter) attached by a pad or knot of fibrils........................................B. echinella i. Fructification larger A. Spores 5-8 t in diameter; carpophore not furnished with a pseudorhiza........................................ B. atrobrunnea A. Spores 4-5 u in diameter; carpophore with a thick well-developed ps udorhiza...........................................B. radicata Bovistella radicata (hMont.) Patouillard (P'ls. XXII-XXIII) Fructification subglobose to top-sllaped, 3-7.5(14.5) cm. broad, smooth to plicate around point of attachment, attached to soil by a thick rooting base 2 cm. or more long, surface over upper flattened part covered with a coating of slender spines, wicl Imay beconle united at tleir tips into compound warts or fascicles, much granular to furfuraceous material present between them, towiard the margin and down the sides the spines become less distinct and finally tlhe surface is merely furfuraceous, lwhite at first, yellowish near maturity, wearing away irregularly to expose the thin papery spore-sac membrane, which eventually qopens by a pore or slit that finally enlarges to expose most of gleba; gleba wlite tlhen yellowish to olive and finally olive brown to dark yellowish brown; sterile base well devel

Page  73 LYCOPERDALES 73 oped, occupying most of narrowed part of Iructification and extending up the sides to form a sort of cup in which the gleba is situated. Spores oval, smooth but with a thin hyaline envelope, 4-5 x3.5-4.5 P in diameter, with a hyaline pedicel 6-11!p long; capillitium of separate units, much branched and branches often quite gnarled, main axis 8-1o p in diameter, ultimate branches tapered to acute or some tapered rather abruptly, filaments often sinuous. Habit, habitat, and distribution.-Potter obtained good specimens of all sizes in the vicinity of Ithaca, Gratiot County. My illustrations are taken from his specimens. Parts of 2 collections by B. O. Longyear are in the Lloyd collections at the Smithsonian Institution. Longyear's material was slightly immature. There are 2 collections at Michigan State College made in 1896 and 1897. The species is apparently common from Ohio southward, to judge from the comments of Johnson as well as those of Coker and Couch. It apparently prefers cultivated fields, though it is also known from pastures and open woods. Discussion.-Coker and Couch reported that when still white clear through the taste is sweetish and pleasant, but that after the color has started to turn a rather strong nitrous odor develops. At first sight one is likely to mistake this fungus for a form of Lycoperdon, but I know of none of this genus in Michigan with such a thick well-developed rooting base. Under the microscope the capillitium distinguishes it at once from all species in Lycoperdon. Bovistella echinella (Pat.) Lloyd Fructification small, 5-8 mm. in diameter, globose, attached by a sandy pad or small knot of debris, white when young and covered by a loose floccose coating, which soon becomes aggregated into floccose patches or gives the surface an almost granular appearance, gradually wearing away to expose the darkbrown papery spore sac, finally nearly glabrous; mouth apical, definite, becoming toothed to lacerated; gleba olivaceous brown at maturity; sterile base absent to slightly developed.

Page  74 74 PU7FFBALLS AND ALLIES IN 1MICHIGAN Spores globose, 5 —6 p in diametecr, withl a tlin liyaline envelope into which project minutte echinulations, olivaceous in KOH, rusty brown in iodine, pedicel 4-8 p long, tapered; capillitium of separate units of long tircads, often septate and more sparsely branched than in Bovista, branches tapered to acute apices, main threads 5-6 p in diameter, dark rusty brown in iodine, duller and more olivaceous in KOH. Habllit, habitat, a(1 distribltionl.- GrCeoarious to scattered on sandy soil in pastures. Potter collected it in (ratiot County, on June 20, 1947, in such a habitat. His collection is the only one I lhave seen, but Lloyd reported it for tle state. Discussion.-In the field tlis funguis is most likely to be confused witll LycoJ)cr)don) p1nsillnl. iotlh are smnall, and the type of flocculen(c over the spore sac is similar enotlugll in both to be confusing. In Potter's specimen the capillitilu, as seen in a cross section of the frluctific(ation, is arranged much as in typical species of Lyco)cr(-lord, but it pulls away from the walls so easily that tcere is certainly little if any attachment. From the material I have seen, B. eC17Jiclla can be distinguished from L. pusilllin lby its larger pedicellate spores. Bovistclla atrobrn? iica Zeller "Fructifications oblate spheroid to turbinate, often collapsed above, 3-4 cm. broad, 2-3 cm. high, with a prominent attachment but not particularly radicate; surface dull, smooth to somewhat futrfulraceous, sonmewhat rimose, dark brown (dry); peridium duplex, exoperidium thin, brittle, breaking up into small plates which easily separate and fall away; endoperidium very thin, papery, dull to shliny, a little lighter colored than the exoperidium, dehiscing by a torn irregular apical pore; sterile base prominent, somnewhat convex above, occupying one-third to one-lalf of the lower portion of the fructification, of large cells, separated by thin, shiny, metallic-brown walls; gleba pulverulent, dark vinaceous brown or darker; capillitium free, long, sliglitly branclled or simple, with long tapering narrow (or even tlread-like) terminals, dark brown, somewlat uneven; spores brown, splirical, verrucose, 5-8 p, with a hyaline pedicel ulp to o30 long wiic:l is easily broken away.

Page  75 LYCOPERDALES 75 "()n tle ground, Ann Arbor, Mlicligan, October 6, 1936, A. H. Smith 5048a-type (in U. Mich. Herb., portion in Zeller Herb.). "Bovistella atrobrunnea differs from other species of the genus in the very dark gleba, the spherical, verrucose spores and particularly in the capillitium. The latter is nearly simple but, now and then, dichotomously branched and the branches or terminals are tapering or drawn out to very long, narrow thread-like filaments very much narrower than the main stem of each unit of capillitium. It is named for the very dark brown gleba" (Zeller, 1948: 649-50). Dicussiou-.-I mistook this fungus for a member of the Lycopclrdon unlbrinum complex, because of its verrucose spores and dark-colored gleba. It may be distinguished from that species by the characters of tle capillitium and the much larger spores. Since it was growing in a conifer plantation, it is not certain that the species is actually native to Michigan. Bovista Persoon Fructification globose to subglobose, often plicate around point of attachment, with a central basal attachment; surface at first covered with a thin, smooth to unpolished covering which flakes off at maturity exposing the smooth polished wall of the spore sac, opening by an apical pore, becoming detached from the ground and then often blown about by the wind; wall of spore sac resilient and persistent; sterile base absent. Spores dark brown to purplish brown and with or without a pedicel depending on the species; capillitium not connected with the wall but composed of separate units which branch irregularly or dichotomously and end in tapered points. The members of this genus, although few, are actually the true puffballs. Coker and Couch treated only 3 species in tlleir work, all of 1which occur in Michigan. KEY TO SPECIES OF Bovista i. Fructification attached to soil by a drab patch of fibers; wall of spore sac at maturity bluish to purplish umber....................B. plunibca i. Not with above combination of characters............................. 2

Page  76 76 PUFFBALLS AND ALLIES IN MIICHIGAN 2. Fructification 3-9 cm. in dialmeer, attached b)) a small cordlike rhizomorph......................................... ila 2. Fructification 1-3.5 cm. in diameter, with a broad basal area of attachm en t.................................................. B. m inor Bovista p)ila Berkeley and Curtis (I'1. XXIv, Fig. i) Fructification -9 cm. in diameter, globose, subglobose or rarely broadly ellipsoid, attached by a small cordlike rhizomorph whiclh breaks easily; surface covered with a thin coating less than.5 Imm. thick which is white and unpolished to furfuraceous and tends to stain pinkislh to alutaceous when handled, becomilng rimose and eventually flaking off to expose the dark brown to bronze, polished wall of tlhe spore sac, usually lhaving a metallic luster, opening at the apex by an irregular fissure whlich becomes enlarged and secondarily torn or lobed: gleba purplish brown. Spores dark brown, globose or nearly so, 3.5-4.2 p, smooth, often w7ithi a broken stub of a pedicel, oil drop eccentric; capillitium dark rusty brown in iodine, dull cinnamon brown in KOH, units miuch branched, the ultimate branellcs pointed and thornlike, main thread 10-12 p in diameter, branches thinner. Habit, haitat, a dit dis lribl ution.-Solitary, scattered to gregarious or sul)cespitose, in pastures, pastured woods, around stables, or where cattle lhave been wintered. 5Morten Lange found it in abundance in an abandoned barn near tle University of Micligan Bioloogical Station at Douglas Lake, Cheboygan County, in June, 1iJ47. Tlle specimens had apparently overwintered, )but were in good condition. Thle fructifications develop dturing tile late sinmmer and fall. It occurs throughout tlhe state and is a relatively common species. Discussion.-Although a typical puffball in tlle strictest sense, one should not expect to find many perfect spheres. The base is often strongly plicate and the point of attachment seated in a shallow depression, and broadly ellipsoid fruiting bodies are not rare. Tllis is by far the largest of tle 3 species of Bovista. Bovista pila can be distinguisled in the field from B. plumbea by the 1muc1 tllinnler sturface coverin and tlhe dark-brown to bronzeo

Page  77 L YCOPERDALES 7 colored spore sac, which is in contrast to the bluish-gray to purplish-umber sac of B. pJluiDbca. In B. pila the spore sac is attaclhed to the substratum by a single cord, whereas in B. plinubca it is attached by a mass of fibers, which usually cause a patch of dirt to adhere at the base. The most reliable distinction between the 2 species, however, is the difference between the spores. Bovista plumbca Persoon (1'1. XXIV, Fig. 2) Fructification 1-3 cm. in diameter, typically deprcssed-lobose, with numerous fibers which cause a patch of dirt 5 mn. or more in diameter to adlere to the base; surface at first covered by a distinct white layer lwhich peels off in sheets over upper part at maturity or dries down to a coating of whitish areolate patches; spore sac thin, smooth, papery but not readily breaking, bluish gray to purplish umber, darker in extreme age, opening at apex by a rather large nearly circular mouth; gleba dark cocoa-colored at maturity. Spores 5-7 x4.5-6 p, oval with long pointed pedicels (-14 p long, with a tlhin hyaline envelope around the spore into whicll minute plugs project to give echinulate appearance noted under an oil-immersion lens; capillitium rusty brown, main tlreads 15-20 p in diameter, branclied, ultimate branches tapered evenly to acute apices. Habit, habitat, and distrib^liion.-Typically, a futngus of open pastures, golf courses, and similar situations, fruiting during tlie summner months and on into the fall. Mains observed it on June 19, 1932, at Blisswood, Emmet County. I have a good collection (Smithl-25803) from Stutsmanville, Emmet County, July 19, 1917, and Kauffman obtained it from a pasture near Ann Arbor, Washtenaw County, (ctober 23, 1926. Potter lhas found it abundant in the vicinity ol Itlhaca, (ratiot County. All collections contained young and freshly matured specimens. It occurs throughout the state. Discussionl.-For a comparison witll B. pila, see discussion of that species. Both B. pila and B. lumbl)ca develop on the surface

Page  78 78 PUFFBALLS ANI) ALLIES IN MICHIGAN of the groulltl, but tlhe tllir(d l-orm, B. iiiilor Morgllan, develops beneath tlhe surface and breaks through at maturity. This character is of little diagnostic value, if only mature or nearly mature specimens are at hland, but tle color of tle wall of mature specimens will at once distinguish B. pluml7bca from B. minor. Bovista min or Morgan (11. XXIV, Fig. 3) Fructification 1-3.5 cm. in diameter, globose to depressedglobose varying to obovoid, often pinched off near the base and plicate around the point of attachment, with a broad basal area of attachment or at times a short-rootilng process extending out of this area and composed of dirt held together by fibrous myceliullm, white when young because of tlle loose m)vclial outer layer which causes mluch trash to ad(here, developing under the surface but soon becoming exposed or more or less exposed ftrom tlhe beginnin (depending o(n lwhether it occurs on hard soil or in loose duff): white outer coating finally matted down to a thin almost membranous covering but pulling apart as expansion takes place and finally at maturity visible as grayish unpolished areas witlh some dirt still adhering; surface dark reddish brown (much as in B. pila), becoming glabrous and polislled only over tlie upper part; spore-sac wall not as rigid as in tlhe otlecr 2 species and at maturity variously dented and wrinkled, opening by an apical pore; gleba distinctly olivaceous b)rown. Spores 4-5 x 3.5-4 P, oval, appearing smooth at ordinary magnifications but under a hig~h power oil-inmmersion lens seen to have a thin hyaline envelope into which project minute echinulations, pale olive brown in KOH, pale rusty brown in iodine, witl long-tapered peliccls up to 15 p; capillitium of branched tinits, tle ultimate l)rancllcs with pointed apices, main apices 10-20 u in diallltter, in K()I dull olivaccous brown, in iodine deep rusty brown, walls much thickened but cell cavity promilent. Habit, habial)t, rand di-tlributioln. —Tlis is generally regarded as a rare spec(ies, lbut on Septltember 28, S q!6, I mlade a collection

Page  79 LYCOPERDALES 79 (Sllitll- 4930) of ablmt a peck of tllh fruiting bodis in a spruce plantation at the University's Saginaw Forest, near Ann Arbor, Wasltenaw County. l)iscjssion.-As the fructifications mature they develop a sweet, almost sickening, but nevertheless aromatic, odor, which is not retained, as in some gill ftngi, after the specimens are dried. Tlis odor along with tle more or less trash-covered mulch-dentcd sulrface of the spore sac allows the fungus to be tquickly identified in tlhe field. The fructifications approacl tlhose of B. p)l1mn ca in size, but tlhe difference in color of the surface of matutre dri-ed specimens allows tlhe two to 1)e quickly distingulisllcd, if old specileni s or herbariumn nmaterial are compared. BROOMI.IACE AE "Fructifications singly or miany on a stroma, Inostly ovoid, hemispleric, or sulbglobose; exoperidiin thin, wholly or partly disintegrated at maturity, endoperidium papery or tllickisl, laid bare at maturity, opening by an apical pore; capillitium present, threads more or less symmetrical, without a (conspicuctous, thick, main filament" (Zeller, 1949: 49). KLY 10 (.IN tNR.A F 1 iR )(tMI \CI.\Fr. i. Fructifiction more or less singly on0 a stro(mla; su.gl)l)obose......./. - sY,,/,/,si/s 1. Fructifications manr' on a stroma and mostly ovoid..................... 2 2. Stroml tllick, stallked, otr co()lui lna;r.l.............................! ctl('jN 2. Stromna sessile, re.supinate, or patellate.......................... i///o 'slis M YCENASTRA CEAE "Fructifications large, subglobose to depressed globose; peridium duplex, exoperidium thick, spongy, smooth or areolate, endoperidium thick and leathery, or tlhin and membranaceous; capillitium branched with short pointed spines; spores spherical to ellipsoid, verr-ucosc" (Zeller,!9,49: 4!). KFY TO (;FNERA OF MYCENASIRACEAE i. Outer peridium smooth, drying thin; inner peridium thick and splitting stellately at maturity..................................ycnast rut i. Outer peridium breaking u) into iyralida(ll scales; inner )peridiuln thin............................................................. C a lb ovij st.

Page  80 80 PUFFBALLS AND ALLIES IN MICHIGAN Mycenaslrum Desvaux Fructifications globose or nearly so, varying to pear-shaped, wall composed of 2 layers, outer layer (exoperidium) thick at first but becoming thinner and floccose, white, becoming broken up into blocklike areas which finally dry down to thin patches and fall away; inner layer (endoperidium) thick (2 mm.+~), indurated and persistent, opening at apex by stellate fissures which gradually extend toward the base dividing spore case into a number of segments which may become recurved; gleba olivaceous becoming umber to purplish brown; sterile base absent. Spores globose, with a thin hyaline envelope almost obscured by the numerous warts; threads of capillitium free from endoperidium, thick, short, branched and covered with thornlike processes. Only 1 species is known from the Great Lakes region. Mlycenastrurn corium (Guers.) Desvaux (P1. XXV and P1. XXVI, Fig. 1) Fructification (4)6-15(20) cm. in diameter, globose, subglobose, obovate, or pyriform, sometimes plicate around the rather broad fibrous mycelioid area of attachment, covered at first by a thick, felted, whitish coat which is continuous at first but soon becomes cracked into blocklike areas and collapses finally to form fairly thin patches of grayish fibrous tissue which eventually fall away exposing the hard to corky nearly smooth surface; surface deep brown to deep purplish brown, the spore-case wall about 2 mm. thick and opening at apex by irregular fissures which become extended toward the base, the segments curve back so as at times to give the appearance of a giant member of Geastrum; gleba white when young, then yellowish olivaceous to olive brown and finally even purplish brown. Spores globose, 8-12 p in diameter, with a thin hyaline envelope almost obscured by the closely set warts which on many spores are fused into an irregular reticulum; capillitium of separate units with the main axes 20-30 [p in diameter, branched freely and ultimate branches as well as main branches (in some)

Page  81 LYCOPERDALES 81 with short spines, walls somewhat thickened and dark rusty brown in iodine. Habit, habitat, and distribution.-Gregarious to scattered in old pastures and fields and around stables where stock has been fed. The material photographed was collected May 31, 1940, but old spore cases from the past season were also present. This is a common species westward in the United States, but apparently fairly rare in Michigan. Most collectors however, find it. Discussion.-This is one of the easiest of all puffballs to recognize at sight in the field or laboratory. The thick wall of the spore case is a reliable field character in old specimens, and the areolate appearance of the immature ones is not possessed by any other species of comparable size in the region. Under the microscope the thorny capillitium and large-warted reticulate spores are distinctive. Coker and Couch pointed out that at maturity it breaks loose from its point of attachment and goes tumbling about as do forms of Bovista. MESOPHELLIACEAE "Fructifications hypogeous or epigeous, singly or several in a stroma; peridium usually 2-3-layered, indehiscent or rupturing irregularly at the apex; capillitium unbranched; spores globose or ellipsoid, variously roughened or with a gelatinous sheath" (Zeller, 1949: 50). KEY TO GENERA OF MESOPHELLIACEAE i. Spores globose and echinulate, reticulate or verrucosc................. 2 1. Spores ellipsoid, smooth or irregularly roughened.................... 4 2. Gleba with a sterile base....................................... Radiigera 2. Gleba without a sterile base......................................... 3 3. Endoperidium opening by irregular ruptures......................Abstomna 3. Endoperidium opening by a simple pore or lacerated opening.... Bovistina 4. Gleba with a large central core........................................Mesophellia 4. Gleba without a central core.................................... Castoreum GEASTRACEAE "Fructifications at first hypogeous, or epigeous from the first, rounded or stalk-like below; peridium duplex; outer peridiull 2-3-layered, pseudoparenchymatous layer within surrounded bv

Page  82 82 PUFFBALLS AND ALLIES IN MICHIGAN a fibrous layer, at maturity opening out stellately (in Trichaster the inner and outer peridium opening togrether); inner peridium papery thin, loosening from the outer peridium and dehiscing by a pore, or irregularly, or by many pores; gleba witl a sterile columella from whicl the tubular cihambers radiate" (Zeller, 1949: 50). KIY 10 (.ENERA OF (CEASTRACLAE i. Endoperidium exposed as a whole at maturity, opening by one or several p ores............................................................ 2 1. Endopcridium disintegrating at maturity.............................. 2. Endoperidium sessile or on a short stalk, opening typically by a single pore (see Astracus also)................................... Gcastrnn 2. Endloperidium on several slender stalks; opening with several pores............................................................... I.l riosto nm a 3. Endopcridium with a prominent sterile base; columiella soft, weak. Terrosiclla E. Endoperidium and exoperidium remaining joined and opening together stellately; columella lard and subwoody; sterile base lacking.... Trichaster MIyriosftoia Dcsvaux Fructification globose to sbl)globose, subterranean until maturity, outer layer coriaceous, pliant, splitting stellately with the segments expanded or reflexed; spore sac witl a membranous wall opening by several pores instead of just one, supported by several short stalks; columellae several, slender; tlireads of capillititun free, unbranclhed. MyIriostoin a coliforme (Pers.) Corda Fructification at first subterranean and subglobose, 1.5-7(10) cmi. broad wlhcn expanded; outer layer nearly smooth or witl adlhering soil particles and at maturity splitting as in Gcastrum into 5 to 7 acute lobes, the fleshy layer thin; spore sac subglobose to compressed, minutely rouglhened, silvery brown, opening by sevcral to many small mouths, sulpported on several short stalks. Spores distinctly warty, 4-6 p; capillitium of long slender tapering tlireads 2-5 p in diameter, witlh thickened walls. Habit, habifat, and distrib titio. —An excellent collection of tllis species collected by C. F. Wheeler, September 2, 1898, at Sandusky, Ollio, is in tlhe MIichigan State College Herbarium.

Page  83 LYCOPERDALES 83 I have seen no material that las actually been collected in Michig(an, but it is likely that tle plant will eventually be discovered in at least the southern tier of counties. Discuissiolo.-Tllis is a rare fungus with an erratic pattern of distrilution. One is tempted to consider it as a form of Gcasir171m, in whicll a nunmber of fruiting bodies have become fused. It is interesting to note that Kambly and Lee reported it from Iowa on the basis of a collection made there in September, 1891. Gcast rum Persoon Fructification subglobose to globose but acuminate at apex, occurring in tlle soil or on the surface; outer wall of 3 layers (an external mycelial layer, a middle fibrillose one, and an inner fleshy layer which at first closely surrounds the spore sac, but is distinct from it), splitting from the apex downward into a numbler of segments (rays) which may become revolute or remain involute; spore sac sessile or pedicellate, melmbranous to papery, thin, globose to elliptic, smooth or roughened; opening by a single apical pore; gleba composed of capillitium and spores. Spores globose, colored, typically roughl; capillitium of sinple, long, tlick-walled, typically pale (revived in KOH) threads arising from wall of spore sac or pseudocolumella; pseudocolumella present or absent. Tlis is one of tle most highly specialized genera of puffballs from tile standpoint of thle organization of the layers in the outer wall of tlle fructification and their behavior. Cunningham stated that upward ofl 1 o species have been dcscribed but that only aboutt:o are valid. Hec rccorded 23 froml Australia. Tile nulmber of species (12) which have been collected in Michigan is not large. Tllis indicates in a imasure tIle ned for more extensive collecting tthroughlout tile state. KL:Y TO SIL::CILS OF Gca(ls1F iill 1. Rays hygroscopic (expanding whein inoistcncd)........................ 1. R ays not as above................................................ 3 2. Peristome sulcate........................................G.. u ilicutum 2. N ot as above...............................................see A stracus

Page  84 84 PUFFBALLS AND ALLIES IN MICHIGAN 3. Mouth sulcatc-striate to conspicuously longitudinally wrinkled......... 4 3. N ot as above....................................................... 7 4. Spore sac asperulate (rough to the touch); spores 6-7.5 A in diameter............................................................ G. cam pcstre 4. Not as above, spores rarely (6 in diameter........................... 5 5. Spores 4-.5-G6; pediccl of spore sac sulcate at zone of attachment to spore sac............................................... pccltinalum 5. N ot as a o ve....................................................... G. Spore sac elevated ill expanded carlpophorcs, the pedicel distinct but short.................................................... Schl idclii 6. Slpore sac practically sessile and remaining seated in a bowl much as in G. sacrat 1u..........................................G. IMorgalii 7. Typically large, 5-10 cm. when expanded............................ 8 7. Usually not over cm. whn expand(led.............................. 10 8. Fleshy layer of rays thick, but breaking so as to leave spore sac sitting in the saucer-like )ase.......................................G. tri l x 8. Spore sac elevated at mnaturity; fleshy layer not splitting as in alove choice......................................................... 9 9. Mouth area paler than remainder of spore sac; outer layer of carpophore separating. leaving under side of rays clean................G. limbatulm 9. Mouth area not sharply delimited; outer surface of carpophore with much trash adhering to it and not separating.............. G. rufescezns 1o. Mouth area sharply delimited from remainder of spore sac by a circular ridge or depression.............................................. 12 1(. MA outh area not distinct........................................... 11 11. Capillitium and spores dark-colored............G. fimbriatum f. fimbriatumn 11. Capillitium and spores whitish.................. G. fibriatum f. pallidunm 12. Spore sac sessile in a bowllike depression......................G. saccatum 12. S1)ore sac pedicellate, wall roughened with minute particles giving it a hoary sheen............................................ G. coronatuLn Geastrum u mb1ilicalurn Fries sensu M organ (as Geaster unmbilicatus) (1'1. XXXI, Fig. 2) Fructification 1-2 cn. in diameter, subgclobose; exoperidiumr separating into 5 to 7 rays, wilic'l in tlie dried state are ascending witl recurved tips, somewhat hygroscopic, rays gradually spreading ott wiien moist; outer layer mycelial and intergrown with debris causing exterior to be more or less evenly covered with sand particles; in some old specimens the debris and mycelium peeling off as a thin layer; flesly layer 1 mm. or less thick, flragile (not as touglh as in G. aslper), light brown when fresh, more of a tan toward base of rays, becoming "bone brown" (Ridgway) or near it on drying (rather dark); surface wrinkled

Page  85 LYCOPERDALES 85 in drying or nearly smooth; spore sac up to i cm. in diameter, globose, sessile or nearly so, pallid because of a tlin whitisl farinose covering whicl wears away exposing the bone-brown ground color of spore-sac wall; mouth area delimited by a depressed zone and usually army brown to bone brown (contrasting sharply witll pallid condition of spore sac on freslly opened carpophores); peristome conic, sulcate. Spores globose, 3.5-4.5 [, minutely warted, wood brown or darker in KOtI; capillititum tlireads wavy, pallid to avellaneous or near wood brown, with some incrustation. Halit, halitat, and1 distri)l)tioli.-Closely gregarious on sand and debris at base of old pine stump in open mixed woods; collected by Victor Potter (Potter-9310), October 15, 1949, from near Sumner, Gratiot County. Discussion.-Thle description is drawn from Potter's notes and from his collection, wlllicl is ample and in good condition. The species concept used is tlat of American autliors. Gcasltrumn coroinatl in Persoon (1PI. XXVI, Fig. 2 and P1. XXVII, Fig. i) Fructification when unexpanded more or less globose and covered by a dense coating of white mycelium with mucl debris intermingled, 0.5-2 cm. in diameter; outer wall splitting into 4 to 6(8) usually strongly recurved segmenits in which the mycelial layer may either become stripped off and remain as an indistinct cup beneath the fruiting body (fruiting body arched upward and remaining attaclled to cup only by the tips of tile rays) or adhering to under surface of rays (Gcastcr coronalllts f. 'inimilus sensu Coker and Couchl); flesly layer dark axellanceots to wood brown often scaling ilore or less comlpletely from fibrillose layer, central part often more persistent and forming a slight collar around base of the pedicellate spore sac; spore sac variable in sllape from subellipsoid to ovoid or subglobose, pedicel 1-2 mm. long and expanding ablove into a ridge or acutemargined rim in the form of a narrow collar, membrane papyraceous, surface toughelned witil minlute glisteningl particles wllich give it a hoary sleen, ground color dark wood brown to

Page  86 86 PUFFBALLS AND ALLIES IN MICHIGAN purplish drab; mouth distinctly fimbriate from radiating fibers which in older specimens converge at the edge into minute teeth, at first forming a small pointed cone, silky area of mouth outlined by a distinct groove or line and paler tlan remainder of fruiting body; gleba dark purplish brown to clocolate at maturity. Spores dark dull brown in KOH and in iodine near sepia, globose, 3.5-5 P in diameter, covered with hyaline truncate warts whlichl more or less cglatinize in KOH; capillitium of very thickwvalled (lumen of cell visible as a line) tlreads, pale yellowish to yellowish in KOH; only sliglhtly darker in iodine. Ilabit, Ihabitat, and d(list ribution.-This species is particularly abundant in a local spruce and pine plantation in the Saginaw Forest, near Ann Arbor, WNaslltenaw County. I suspect that in tlhe state it is locally abundant in northern regions during wet seasons. Potter has seen it on a number of occasions near Ithaca, Gratiot County. The fructifications are usually scattered to gregarious with only the top of the white mycelial ball visible when immature. The fungus occurs locally and fruits during September and early October, depending on how soon the rains come. Discussion. —llis is a very easy species to recognize because of tle copious whlite mycelial coating of tlhe unopened fructifications, tlhe peculiar slheen over the dark pedicellate spore sac with its basal collar, the fibrillose mouth delimited by a distinct pale circle, and tlhe manner in which the fructification typically stands up on the points of the rays when fully expanded. As is often tlhe case with a (ommon species, this one has been given a number of specific names. For a discussion of these names and tlle variable claracters which caused them to be applied, see Coker and Couch. Gcastlrun pcctiuatum Persoon (1'1. XXVII, Fig. 2) Frtuctification 1-2 (cm. in diameter in unopened condition, outllines obscured by copi)ous amount of trash adhering because of well-developed nlycclial layer, opening by outer wall and splitting into 8 to io rays (in tlhe Micligan collection); fibrous layer recurving until inner surface is more or less convex and

Page  87 LYCOPERDALES 87 outer surface reversed so as to line the concave part, sinuses of rays extending about half way to point of attachment of spore sac; fleshy layer (covering convex surface) near wood brown to cinnamon brown and in drying cracking into very irregular patterns and, according to most authors, flaking off at maturity, much debris held in place in the concave part of expanded fructification; spore sac subglobose to urn-shaped, 1-2 cm. in diameter, purplish drab beneath a hoary sheen; apophysis present or absent, part which narrows to pedicel often radially sulcate, smooth in some, according to most authors; pedicel up to 6 mm. long and often with collar or ring near the middle or base; mouth conic, sulcate-striate, at base not sharply separated from spore-sac membrane; gleba chocolate brown. Spores globose, 4.5-6 p in diameter, bister (dark dull brown), covered with flat-topped hyaline warts; capillitium threads 4-7 p in diameter, in KOH the walls sordid yellowish to brownish, in iodine paler and brighter yellow, very evenly tapered to apices, unbranched to sparsely branched. Habit, habitat, and distribution.-A very rare species generally, known from 5Michigan from a collection made by Hugh Iltis under cedars in the gorge near tle University of Michigan Biological Station at Douglas Lake, Cheboygan County. Two fructifications were found, both in good condition. Discussion.-The sulcate conic peristome, which is not sharply delimited from the spore sac; the sulcate underside of the spore sac, where it joins the relatively long pedicel, and the welldeveloped mycelial layer are the conspicuous field characters in typical material. Gcastrum Schmidclii Vittadini (P1. XXVII, Figs. 3-4) Fructification small, when unopened 5-8 mm. in diameter, globose with a bluntly conic apex, pallid and with adhering trash, opening by outer wall splitting into 4 to 6 rays which spread out into a horizontal or slightly recurved position; mycelial layer persistent causing trash to remain on under side of rays; fibrous layer very thin in dried specimens; fleshy layer also

Page  88 88 PUFFBALLS AND ALLIES IN MICHIGAN thin, continuous out over surface of rays or in drying cracking at juncture of rays and base of fruiting body, pallid to pale brownish; spore case ovate to globose-pedicellate with a long conic peristome; surface pallid to brownish drab (in oldest specimen), papery, unpolished to covered witlh a whitish bloom; mouth strongly sulcate, grading indistinctly into tissue of spore sac or obscurely delimited by a depression; gleba chocolate brown at maturity. Spores 3.5-4.2 p in diameter, globose with a thin hyaline envelope into which slightly colored warts project. near bistcr in color in iodine under the microscope; capillitium pallid to slightly yellowish in iodine, tlreads 3-6 p in diameter witli very thick walls. Habit, habitat, anzd distribution.-On rich humus in hardwood forest, apparently gregarious. Not uncommon in the central part of the state. In the herbarium at Mlichigan State College there is a collection which was made September 25, 1900. Kauffman and a party of students obtained it near lWhitmore Lake, Washtenaw County, October 5, 1923. Potter collected it rather frequently in the vicinity of Carson City, MIontcalm County, in October and November, 1948. Disculssioin.-This species is at once distinguished in the field by tle strongly sulcate peristome, by the slort pedicel of the spore sac, and by its not being fornicate when fully expanded. Geastrum Schnlidclii is closely related to G. pectilaltunl but differs in not being striate wlhere the spore sac tapers to the pedicel, in the slorter pedicel, smaller spores, and in the rays not becoming bent so far back. Gcastlrnn Alorgantii Lloyd (1'1. XXXI, Fig. 3) Fructification b1roadly bulb-slaped when unexpanded, 1-2.5 cmn. broad, point of attachment central, base flattish, outer surface vinaceous buff to wood brown (witll a vinaceous cast), witll some dirt and trash adlhering but generally fairly clean, tile layer splitting sonmewiat on drying and with a tendency to separate in small areas exposing thle pallid smooth shining under

Page  89 LYCOPERDALES surface; inner surface (fleshy layer) about 3 mmin. thick, soft, watery, pallid to pale pinkish (resembling that of G. rufescens), and turning rufous when bruised, gradually becoming reddish brown when dried, finally near wood brown and thin, variously cracked or forming a collar around spore sac; spore sac 1 cm. broad, subglobose when dried, wood brown to near bone brown, surface felted and unpolished; mouth area delimited by a circular line but concolorous with rest of sac; peristome conic, longitudinally wrinkled to sulcate. Spores 3-4 (4.5) P,1 globose, minutely warty, near bone brown in mass, near bister under the microscope; capillitium threads 4-8 p in diameter, wavy, hyaline to dingy yellowish brown, thick-walled. Habit, habitat, and distribution.-Gregarious around hardwood stumps, Itlaca, Gratiot County, September and October. Disclussion. —This species is known in Michigan from 4 collections by Potter in the vicinity of Ithaca and 1 by Harper from Saugatuck. It resembles G. saccatum in general appearance but is distinct by virtue of the more or less sulcate-striate peristome. Geastrum triplex Junghuhn (I'1. XXVIII) Fructification typically large, 1-5 cm. in diameter in unexpanded condition, up to 8-9 cm. broad in some as rays spread out, witll the apex produced into a pointed beak 6-18 mm. long (measurement made on dried material), typically sessile, in large specimens the base somewhat gnarled occasionally, grayish brown to wood brown when young, dull ochraceous to alutaceus in dried (unopened) specimens; outer wall splitting into (4) 5 to 7(8) rays (primary rays usually 4 in number, splitting secondarily to produce tile larger numbers mentioned above); outer surface of rays and unopened specimens rou(gh but witl very little trash or dirt adhering; base marked by a distinct scar at point of attachment; fleshy layer of rays (inner layer) near wood brown wihen dried, often separating from rays near their bases as a continuous sheet thus forming a cup in which the

Page  90 go PUFFBALLS AND ALLIES IN MICHIGAN spore case is seated, in some specimens remaining attached as an adnate sheet over unsegmented part of outer wall, the part adhering to the rays variously cracked and sometimes finally peeling off in places; spore case sessile, pale to dark avellaneous, unpolished, mouth conic, delimited by a circular paler area, distinctly radially fibrillose, the fibrils at the apex in groups of unequal length to form a more or less lacerate-fibrillose mouth. Spores globose, 3.5-4.5 p in diameter, covered with short narrow truncate projections (under a high power oil-immersion lens) of a hyaline substance, pale cinnamon brown in KOH, dark dull brown (near sepia in iodine); capillitium of more or less encrusted filaments 3-6 p in diameter, the walls thickened to the point where the lumen of the cell appears as only a line (hence one might mistakenly think the cells were thin-walled because the lumen cannot be readily seen), color varying from hyaline to dull yellowish brown in KOH, merely yellowish in iodine. Habit, habitat, and distribution.-Single to gregarious in hlardwood forests in which much humus has accumulated. The fructifications are at first almost buried in the loose duff but emerge at maturity. It is the commonest form of Geastrum in the hardwood forests of the state. Good collections of unopened specimens may be made during August and early September. After that the expanded specimens are the ones most frequently encountered, and the old spore cases may survive the winter and be picked up the following spring or summer. Discussion.-The distinguishing characters are the pronounced beak of the immature fruiting bodies, the lack of trash adhering to the outer surface, the saucer-like to cuplike depression in which the spore case is seated, the relatively large size, the fibrillose mouth, and paler surrounding area more or less distinctly setting it off from the remainder of the spore case. Geastrum limbatum sensu Coker and Couch (as Gcaster linmbatus sensu Bresadola) (1'1. XXXII, Figs. 2-3) Fructification typically large, 2-4 cm. in unexpanded condition and up to 6 cm. or more across the tips of the rays when

Page  91 LYCOPERDALES 91 expanded; outer peridial surface drying hard and firm, practically clean of trash, upon expansion of the fructification separating cleanly from medial layer over the central part and almost to the tips of the rays, thus leaving a cup in the ground to which the tips of the rays are attached in the expanded fruiting body; fleshy layer (inner or upper layer) dull cinnamon when dried and irregularly to radially rimose, separating slightly along margin of rays; spore sac subglobose, dark wood brown to light drab, short-pedicellate as seen in expanded carpophores; pedicel pallid; mouth area pallid and fibrillose. Spores 3.5-4.5 p in diameter, globose, warty, pale bister revived in KOH; capillitium of hyaline to dull brown thick-walled filaments, smooth or with some adhering debris. Habit, habitat, anld distrib ution.-Thc only collection from the state I have recognized is one by Potter (Potter-8770) from Crystal, Montcalm County, taken September 14, 1949. It was on the surface of the ground at the base of a fence post at the edge of a sandy pasture. Discussion.-The definite mouth area distinguishes this species from G. fornicatum. From G. rufescenls it can be distinguished by the lack of vinaccous colors and by the way the outer layer separates as the rays expand. The concept of the species as I have used it here is not that of Cunningham. H-Ie suggested that the concept of Coker and Couch, the one I have followed, really applies to a subpedicellate form of G. triplex. No such assignment of Potter's collection is reasonable. Geastrum rufcscens Pcrsoon (1'1. XXIX) Fructification at first flattened to globose and apex pinched off somewhat to form an obscure beak, some merely convexovate above a flattened base, buttons (dried) up to 3.5 cm. broad, more or less buried in the duff and witli a layer of debris held to the exterior by a thin separable mycclial layer which peels off exposing the avellaneous to pale fawn color (vinaceous) outer surface of the cortex, cortex thick, coriaceous, splitting about halfway to base into 5 to 9 lobes with fairly acute apices,

Page  92 92 PUFFBALLS AND ALLIES IN MIICHIGAN primary ]l)obcs oc(casionally split into seconlllary lobes lbit tle latter usually short, the fleshy layer tlick (5 mIm. +) when fresl, forming a thin continuous crust or l)reaking uip somewhat as in G. triplcx; spore case sul)globose, 1-4 cm. in diameter; substipitate, tle stalk rather thick, sometimes with a low apophysis some distance fromn stem on lower side of spore sac, surface of spore sac dull brownish and unpolished to granular-velvety; imoutlh slightly elevated, fibrous, and becoming finlbriate; witlhout a definite peristom)c'; colunll llla g(lobose to subglobose; gleba lister or darker. Spores (,'lobose to sulbglobose, 3-4.2 p in diameter, minutely warted, with a thin gelatinous envelope; capillitiumi of simple tr-ca(ls 3.5-( p in diameter, thick-walled and walls smooth to incrusted, pallid to dull brown in KOH. Habit, 1ihal)ita, and(( d(islribu)iol.-C(cspitosc-grecarious in d(lCris on light soil around hardwood stumps. Probably throughout tile state. Potter collected it abundantly in the vicinity of Itlaca, Gratiot County, in Augusit and September, 1948. Discussion.-The dlescription and illustration are taken from Potter's collection. Coker and Couch reported a fruity odor to tihe fresl specimens. Although Gcastru n r^-fesceis is a large species on tile order of G. triplex it can be readily distinguished froml the latter by the vinaceous color of the fibrous layer once tle debris is remo(vecd,, by tile spore case becoming elevated on a short stalk, and by the fact that tlhe cortcx in splitting becomes arcihed snuffcicn tl to c'lcvate tlhe spore sac (compare Plates XXVIII-XXIX). Gcastrumt firn7lriatum Fries f. finmbriatuml Fructification typically small, o.8-2 cm. in diameter before opening, depressed-globose witl a mucronate apex in dried specimen (sce Discussion), the outer nycelial layer pallid to buff (cinnamon buff), 'formlingl a tliin fibrillose coating; outer wall splitting into 5 o 8(12) rays wlhiclh have acute tips, sinuses extending dow-n for alout half the diameter of the fructification leaving tIle spore sac nestled in a bowllike base, the rays finally recurved back to almost toucling tle exterior of the bowl; my

Page  93 LYCOPERDALES 93) celial layer tending to pcl or flake off leaving the smooth pallid fibrous layer exposed; fleshy layer mucl as in G. saccatuilm, when dried a thin, tan to dull brown coating over bowl, tending to become rimose at juncture with rays and on their surfaces; spore sac globose with a conic mouth, 8-15 mm. in diameter, unpolished, tan to dark brown; mouth fibrillose, not distinctly marked off from rest of spore sac but in some witl a slightly different color or sheen; gleba fuscous brown. Spores small, 3-3.5 1 in diameter, glol)ose, with a very tliin hyaline envelope and minute projections extending into it, dark brown (near bistcr); capillitium threads nearly hyaline (both in KOH and in iodine), very thick-walled, unbranched, 4-7 [I in diameter. Habit, habitat, anld distrib)tion.-Scattered on rich humus in low woods of deciduous trees. I have it from 2 collections, one at WXaterloo, Jackson County, and the otlier at Dexter, Washtenaw County. Among Kauffman's collections are 2, one from \\Whitmore Lake and the other from Ann Arbor, both in Washtcnaw County. The collecting dates are August 8 to November 2. Potter lhas a number of collections from near Ithaca, Gratiot County. It apparently is a rarely collected species. The buttons develop in the trash, some of which adheres to the mycelial layer as the rays expand. Discussiont.-The mucro on the single unopened specimen of Gcastrum firlbriatlllu collected is either a variation from tile usual or the specimen is a stray button of anotlcer species. I am inclined to believe it a variation, as all other characters check for the species, including the mouth claracters of the spore sac. For purposes of field identification G. fimbriatum micght be regarded as an example of G. saccaturm, without a line delimiting the mouth of the spore sac. There are otlher claracters, such as the smaller spores which distinguish it. Since the mycelial layer over the rays also has sonme tendency to chip off in G. saccatum, not nmuch emphasis las been given this character here. Kauffman considered G. fizmbrialum a synonym of G. saccatum and filed his collections under saccatunm.

Page  94 94 PUFFBALLS AND ALLIES IN MICHIGAN Geastrum fimbriatum f. pallidum, f. nov. A typo differt: sporae hyalinae et capillitio hyalino. Fructification 10-15(20) mm. broad, 10-20 mm. high including the short point, expanding to 2.5-4 cm. as rays open out, outer surface covered by a distinct mycelial layer binding much trash to outer surface, surface beneath trash tilleul buff to near pale cinnamon buff, rays 5 to io, spreading to revolute, or curved in against the bowl-shaped base; inner (fleshy) layer pinkish buff or paler and not darkening in drying, thin and forming an even, continuous coating over the exposed surface of the rays and bowl, in drying remaining smooth or cracking slightly along edge of bowl and pulling away slightly from the margin of the rays; spore sac sessile, 10-15 mm. in diameter, subglobosc, wall papery, pale pinkish buff to whitish when dried and roughened with small white granules; peristome conic, whitish, fibrillose, not sharply delimited from remainder of sac; gleba tilleul buff (whitish). Spores hyaline under the microscope, 2-3.5 (4) p in diameter, very minutely echinulate; capillitium of hyaline thick-walled filaments with occasional cross walls, mostly unbranched, smooth, straight or slightly curved. Habit, habitat, and distribution.-Gregarious on rich humus, Tahquamenon Falls State Park, Luce County, September 2, 1940. Discussion.-I have been unable to find a satisfactory description of this fungus in the literature, and since it is known only from a single collection, prefer not to apply a binomial to it at the species level. It is an albino or near albino G. fimbriatum with an asperulate spore sac. The hyaline spores and hyaline capillitium are curious indeed. Lloyd (Mycological Writings, 1:202, note 304) reported on an albino form of Geastrum triplex sent to him from Canada. It was like the typical form, he stated, in all respects save for the pale spore sac and the hyaline spores and capillitium. Further collections are needed to determine whether it is more than an isolated chance occurrence. Cunningham in the Gastronycetes of Australia and New Zealand (1942: 169) stated, of G. limbatum, that "occasional speci

Page  95 LYCOPERDALES 95 mens have the endoperidium covered with a white farinose coating." This is apparently the same type of coating as that of the Tahquamenon collection. Gcastrum saccatum Fries (I1. XXX) Fructification 6-12(20) mm. broad, 8-15 mm. high including the point which is 1-2 or more mm. long, expanding to 2-5 cm. as rays open out; outer surface with very little adhering trash, ochraceous buff to cinnamon buff, spongy-felted, when dried irregularly rimose and in some pulling off over considerable areas; typically with a basal scar; rays 4 to io, (often split into 4 to 6 primary rays, some of which in turn may be split), revolute to curved in against the bowl-shaped base; inner (fleshy) layer pallid to avellaneous when fresh, drying to near wood brown, thin and forming an even continuous coating over the exposed surface of the rays and bowl, in drying remaining smooth or variously rimose along edge of bowl and on the rays; spore case sessile, 0.5-2 cm. in diameter, globose to subglobose, wall papery, purplish drab to wood brown, unpolished; peristome conic, fibrillose, delimited by a circular raised to depressed line, often paler or more brownish than spore-case wall; gleba coffee color to paler; pseudocolumella well developed. Spores 3.5-4.5 P in diameter (many poorly formed spores in mounts measuring 2.5-3.5 p), globose, pale date brown in KOH and iodine, verrucose with truncate hyaline to yellowish warts; capillitium of pale yellowish to brownish, very thick-walled threads with much debris incrusting them, 4-8 P in diameter, often quite crooked. Habit, habitat, and distribution.-Gregarious on rich humus, often around old stumps, during late summer. Both of my collections were made in August. The distribution in the state still remains to be established, since the species was confused previously with G. fimbriatum. The illustration is of a collection by Potter. Discussion.-In G. fimbriatum the mouth is not sharply delimited by a circular line or groove. This is the important dif

Page  96 96 PUFFBALLS AND ALLIES IN MICHIGAN ference between it and G. saccatur. The rays of specimens of saccatum in my collections have exhibited no distinct tendency to split into 2 layers, but in i collection (Smith-15192) the mycelial layer has become rimose in drying and has flaked off from around the base of the oldest specimen. The Michigan collections, on the basis of spore size, agree better with the typical form of G. saccatun as described by Coker and Couch, than with their northern form. Cunningham used the small spores, 3-3.5 p, as the key character of the species. In some mounts made from the Michigan collections many small spores were present, but they were pale and otherwise appeared abnormal, so that I do not attach any taxonomic significance to them. It is generally admitted that the species is extremely variable, and my limited observations support this belief. The fungus is clearly in the G. triplex group, but is readily distinguished by the manner in which the fleshy layer behaves as the rays spread out. Gcastrum campestre (Morg.) Kambly and Lee (P1. XXXI, Fig. 1) Fructification small, globose, submerged at first but becoming lifted up as it expands, 2.5-4 cm. across when expanded; outer layer splitting past the middle or two-thirds of the distance to tlie base, becoming divided into 6 to 12 rays which usually have a layer of sand held to the under surface by a thin layer of mycelium, the rays expanded when wet and involute when dry, folding over or under the spore case in drying, fleshy layer clocolate color to umber, and soon wrinkled-rimose; spore case sessile to short-pedicellate, 8-15 mim. in diameter, subglobose, color dingy but surface typically ashy gray from a coating of adnate asperulae causing the surface to feel rough; peristome prominent, conic, acute, seated on a more or less distinctly delimited zone, strongly sulcate and often dark-colored; gleba chocolate color. Spores globose, 6-7.5 p in diameter, very dark brown in KOH, coarsely asperulate with hyaline plugs extending into a gelatinous matrix, apedicellate or pedicel present only as a broken

Page  97 SCLEROi)ERMA TALES 97 stump; capillitium of threads 3-5 P in diameter; pale yellowish in KOH, walls thickened, slightly flexuous, rarely branched, no cross walls seen. Habit, habitat, and distribution.-Gregarious in sandy pastures during October and November, near Riverdale, Gratiot County. Several collections from near Riverdale by Potter constitute the only Michigan collections located in the course of this study. Discussiou. —This species is reported under the name G. asper in much of the older literature. The asperulate surface of the spore sac and sulcate peristome readily distinguish it from other Michigan species, but Cunningham in Australia reported a series of 4 species which have the above combination of characters and stated that they intergrade. SCLERODERMAATALES "Fructifications mostly epigeous; sporocarp sessile, on a false stem, or if stipitate, borne entirely above the stem or its expanded summit; peridium of 1-4 layers, dehiscing by an apical stoma, or by irregular fissuring or circumscissilely; gleba pulverulent at maturity, with or without capillitium; basidia symmetrically distributed or in nests or cavities arising through the dissolution of the tissue, without a well organized hymenium (except possibly in Battarrea)" (Zeller, 1949: 51). KEY TO FAMILIES OF SCLEROI)ERMATALES i. Peridium simple (no membranous inner layer present)................. 1. Peridium with distinct endoperidium and exopcridiun................ 5 2. Capillitium well developed; gleba entirely homogeneous.......................G......O A..................................... G IIRO L MAACLAL 2. Capillitium wanting or rudimentary.................................. 3 3. Gleba breaking up into small peridioles...................... PISOLITIACLAE 3. N ot as above........................................................ 4 4. Gleba mostly veined or lacunate, veins developing internally or centrifugally.............................................SCLERODERMAT^ACEAI: 4. Gleba with veins definitely extending centripetally from the peridium-............................................................ S:1):C L'LACI':A E 5. Fructification with a firm or gelatinous stalk.......................... 6 5. Fructification sessile; exopcridium splitting as in Gc(strumln...... AsRAEFCI.AC.A 6. Stalk firm, fibrous, or woo(y.............................. 1UO)SIOMATACEAE 6. Stalk gelatinous fresh, consisting of anastomosed strands to form a rough, lacunose structure.................................. CALOSTOMATACEAL

Page  98 98 PUFFBALLS AND ALLIES IN MICHIGAN SCLERODERIATACEAE "Fructifications mostly epigeous, rarely hypogeous or emergent, subglobose, sessile or with an irregular root-like stem; peridium mostly simple, rarely 2-layered, firm, rarely thin, membranous, breaking open irregularly or in lobes or decaying; gleba with sharply defined basidia-bearing sectors, which are partitioned from one another by sterile veins, and in which the basidia are regularly scattered through the tissue (rarely, if ever, witli hymenium-lined cavities or with fascicled, nested basidia); basidia broadly club-slhaped; gleba crumbling to a powder of spores and disintegrating tissues at maturity; spores usually sculptured; capillitiun wanlting, or rudimentary" (Zeller, 1949: 51). KLY '10 (,INlR.\ OF SCI.ROD)lRMAITA(I:.I 1. Fructification with slender stein................................. Pirogastcr 1. Fructification not as above........................................... 2 2. Peridium covered with spines or coarse Iy)ramlidal warts.......... Cloderma 2. Peridium smooth to finely warted or surface rough (but not with spines or prom inent raised warts)......................................... 3 3. Spores lacking a pediccl and a distinct Iilum.................. Sclcroderma 3. Spores with a hiluln or pedicellate.............................Pomlpholyx Sclcrodcrmna Persoon Fructification globose to pear-shaped or subturbinate, partly to entirely above ground at maturity, often with a false stipe composed of coarse mycelium and dirt; spore case with a thick wall and the outer surface varying from smooth to variously sculptured depending on the species, opening in an irregular manner; gleba composed of tramal plates enclosing cavities, breaking down at maturity to a pulverulent mass and then usually very dark (olivaceous, fuscous to purplish black); capillitium typically lacking; spores globose, ornamented. This is a genus that all collectors soon encounter in the state. Cunningham pointed out that over 6o species have been described, but lie thought that only about a dozen are valid. This is doubtlless too conservative a figure. Zeller (1947) recently described 3 from the western states. Kambly and Lee (1936)

Page  99 SCLERODERMA TALES 99 recognized 3 species from Iowa; Johnson (1929), 6 from Ohio; and Coker and Couch 6 from the eastern United States and Canada. Cunningham recognized only 6 from Australia and New Zealand. KEY TO MICHI(GAN SIPECIES OF Scleroderma i. Fructification with distinct arcolate markings. Spores reticulated.S. aurantiumn 1. Not with above combination of characters............................ 2 2. Fructification typically smooth when young, opening somewhat as in Gcastrum................................................... flavidu 2. Fructification very coarse and much roughened, opening somewhat as in Geastrum................................................ Gcastcr 2. N ot as above....................................................... 3 3. Spores reticulate; fructification 3-9 cm. in diameter.............S. arcricola 3. Spores strongly echinulate; fructification 1.5-4(5) cm. in diameter.............................................. S. lycoperdoides Scleroderma auranttiurn Persoon (PI. XXXII, Fig. 1; PI'. XXXIII) Fructification (2)3-10(12) cm. in diameter, globose, subglobose, or considerably depressed at maturity, sides and under side often plicate or in some actually lobed, sessile, or pseudostipitate (with a stemlike base composed of compactly interwoven mycelium and debris); spore sac ochraceous to clay color (some shade of yellowish brown); surface cracked into distinct areolations which often have a central wart, the warts and areolations often arranged in a beautifully embossed pattern; wall of fructification about 2 mm. thick when fresh, about 1 mm. or less dried, turning pink when fresh material is cut, opening at apex by slowly cracking into irregular lobes whlich do not open out in a stellate manner; gleba gray witli whitish lines of the tramal plates (these olivaceous in some dried material), finally dark violaceous gray to nearly black, with the consistency of a rather coarse powder at maturity. Spores (8)9-12(13) p in diameter including projections; globose, verrucose-reticulate with more or less confliuent warts of a softer substance than the thick, dark-brown wall. Habit, habitat, a(11 distrib uti'on.-Solitary, scattered, gregarious, or even cespitose around old stumps and logs, in low

Page  100 100 PUFFBALLS AND ALLIES IN MICHIGAN rich woodlands, on hummocks in bogs and similar places; in either hardwood or conifer forests, but frequent around hemlock in the nortl and around the large blueberry (Vaccinium co ry mbosunm) in the south. Discussion.-In the field the characteristic pattern of warts, present at least over the upper surface, will distinguislh the species. Under the microscope the large somewhat reticulated spores are distinctive. The extent to whicl the stemlike base is present is a thoroughly unreliable character. The amount of its development appears to be determined by tlhe porosity of the substratum. It is niost likely to be nearly absent on a hard substrattun and wvell developed on loose llumuis or sandy soil. Under a good oil-inmmersion lens tlhe reticulation of the spores is seen to be incomplete; there are a fair number of unconnected warts on each spore. Although I lhave never recommended this species to anyone as a food, it must be admitted that it is eaten by some people. In England it lhas been used as food under the name "vegetable tripe" and also as an adulterant of sausage. Mcllvaine and Macadami reported the North American species, as far as they tried tlenm, edible but that they become bitter and unpalatable (inedible) with the first trace of yellow in the gleba. Tlhe fructifications of this species are sometimes parasitized by Bolclus parasiticus. The form illustrated on Plate XXXIII hlas usually been referred to S. aunranlilulm but I have included it here with great reservations since the markings of the peridium are not typical. Sclcrodcrima flavidu i Ellis and Everhart (11. XXXI7V) irltictificatioln 2-(;(8) (cn. in diamncter,, lobose, becomling subgloblose to deprcssed-globose, the upper surface flattened somewhat, plicate to irregularly wrinkled toward point of attachment, with a tlick fibrous rooting base containing much soil, its area of attaclhment 1-2 cm. or more broad, young specimens almost buried in tlhe sand but more or less exposed by maturity; spore-sac wall pallid yellowish to straw color; upper surface nearly smootl at first but soon becoming3f rimnose-areolate, tlle

Page  101 SCLERODERMA TALES 101 areolations flat (like cracks in the mud) and paler (sordid whitish to dingy gray), along the marginal area sometimes more or less imbricate scaly, surface nearly smooth on under side; wall tough and opening apically by splitting into a number of lobes which may in turn split and which all open outward and recurve somewhat as in Geastrium, at maturity or in age splitting halfway or more toward point of attachment; gleba powdery, pale dull yellowislh brown to very dark yellowish brown at maturity. Spores globose 9-13(14) j in diameter including spines, densely covered with conic spines 1-1.5 P long which appear nearly liyaline at the tips and tend to collapse or gelatinize somewhat; numerous hyaline filaments (from trama) also present in mounts, these are branched and measure 2.5-4.5 in diameter. Habit, habitat, and distribution.-Scattered to gregarious, almost buried in the soil at first. It is a frequently encountered fungus on rather barren sandy hillsides in northwestern Washtenaw County, particularly in the vicinity of Silver Lake and the George Reserve, near Pinckney, Livingston County. Potter has also collected it abundantly. Cunningham reported it as the most frequent species of Scleroderma in Australia. It matures during the late summer and fall. Discussion-.-In Michigan its habitat in sandy soil is a fairly good field character. The pallid to grayish flat scales and the manner in which the wall of the spore sac splits into lobes and then recurves, Geastrum-like, are the important diagnostic points. Microscopically, the strongly eclinulate spores are iunportant along witl the llyaline threads left by the breaking ldown of the trainal plates. Tile old open spore sacs are very persistent and are found intact after overwintering. Sclerodcrma arciiicola Zeller (P1. XXXV, Fig. 1) Fructification 3-9 cm. in diameter, subglobose, urceolate, or cushion-shaped (when mucl broader than thick), sometimes quite irregular or with lobes or deep wrinkles, often stlcate around point of attachment, sessile or with the usual pseudo

Page  102 102 PUFFBALLS AND ALLIES IN MICHIGAN stem made up of a fibrous mass holding much dirt; surface smooth to matted fibrillose but developing minute flat spotlike scales or in some surface areolate-rimose and rather deeply cracked, straw color to pale leather color, finally duller brown but not blackish; spore-case wall thick and firm, very hard when dry, finally opening by a slit or irregular rupture; gleba dark yellowish brown to fuscous or blackish. Spores near mummy brown (Ridgway) under microscope when mounted in iodine (very dark almost blackish brown), globose, (10)15-20(22) pin diameter including reticulation; echinulations also present and up to 3 X long, the reticulum and echinulations of a paler more gelatinous substance than the thick dark-brown spore wall. Habit, habitat, and distributionl.-Scattered to gregarious under conifers or hardwoods on poor soil. I have a number of collections from Washtenaw County and one from Grouse Haven, property of the State Department of Conservation on tlhe Rifle River near Lupton, Ogemaw County. There is a fine collection at Michigan State College, East Lansing, Ingham County, made on the campus September 16, 1896, but filed as S. vulgare Fr. One should look for the species from August to October. Discussion. —Tlis species is listed under the name Scleroderma Bovista sensu Bresadola in Coker and Couch. Zeller has formally recognized it as a species distinct from the true S. Bovista of Fries, S. arcnicola can be distinguished at once from S. lycoperdoides by its tllick hard wall, much larger spores, and lack of a true stipe. The scales of tle 2 species, however, may be very similar. Sclcroclerma lycoperdoides Schweinitz (1'. xXXV, Fig. 2) Fructification (1)1.5-4(5) broad, depressed-globose (broader tlan tlick) to somewhat irregular, stipitate; stipe enlarged upward from a filbrots-mycelioid attachment which may be enlarged into a pseudostipe, apex and under side of spore sac often sulcate; spore case light brown to yellowish brown and

Page  103 SCLERODERMA TALES 103 dotted all over (but more conspicuously on upper surface) with distinct inherent, darker brown scales, in some specimens scales dotlike, but in others flat and subcircular to obscurely rectangular in outline and crowded to widely separated; surface nearly smooth to somewhat roughened from scales, in some specimens the scales sunken slightly when dried leaving surrounding surface in the pattern of a paler reticulum; wall of spore case thin and when dried fragile, inner side not clearly distinct from gleba, finally opening by an irregular slit or fracture (or by insect damage) on upper surface; gleba watery cream color, changing to deep brown and then purplish, and finally grayish to dull olive brown in age, spores intermixed with threads and partly broken down tramal plates. Spores globose, (8)10-15(18) in diameter, variable in size, strongly echinulate, dark yellowish brown in iodine; the spines 1-2 p long; filamentous hyaline hyphae and much partly disintegrated material also present in mounts. Habit, habitat, and distribution.-Gregarious to subcespitose or occasionally scattered, on humus or along very decayed logs, often along the edges of bogs, and in an Ann Arbor arboretum on the mulch around various rhododendrons. In the southeastern part of the state it is often common from the first part of August on into fall, but it appears to be rare northward. A collection of Potter's was taken from bare soil at the bottom of a ditch near Ithaca, Gratiot County. Discussion.-In extreme age only the stipe and a saucer-like expansion representing the base of the spore case are left, and these may persist over winter. The stem, the small size of the spore case, and the small spotlike scales are the important field characters. Scleroderma Geaster Fries (P1. XXXVI) Fructification 4-12(13.5) cm. broad, opening up to 15 cm. or more; globose to depressed-globose, wall thick (5 mm. ), hard, rough, and soon variously cracked, much dirt usually adhering; in section whitish to yellowish or finally clay color, splitting into

Page  104 104 PUFFBALLS AND ALLIES IN MICHIGAN distinct segments at maturity which open out in the manner of Geastrum, to expose the dark powdery gleba; gleba brownish to finally umber, some hyphae (remains of tramal plates?) scattered through it. Spores globose to subglobose, 6.5-9 P in diameter, with a hyaline reticulate sheath and small verrucose projections extending into it, hyaline material slougliing off readily and littering the mount. Habit, habitat, and distribution. —The photograph and description are from Virginia material collected by Kauffman. Coker and Couch reported it as occurring on the sides of gullies in clay soil. There are no records from Michigan, but it should be sought for on the clay soils of the southern counties. Discussion.-Cunningham reported Sclcroderma Geaster from Australia but with spores 14-18 p in diameter; he also pointed out that most previous reports of it were on material of other species. On the basis of the difference in spore size it seems doubtful to me that the specimens admitted by Cunningham can be conspecific with those having small spores from eastern North America. Longyear reported it along with S. flavidumn from Michigan, but the specimens now preserved in the herbarium at Michigan State College, are actually S. flavidumr. Barnett first noted this fact, and my observations support his conclusion. Mains's report, formed on an identification by Kanouse, is also based on S. flavidunm. SEDECULACEAE "Fructifications leatlery, without sterile base or radicle; peridium tlick, leathery above, almost obsolete and dehiscing below; gleba becoming powdery at maturity, with broad veins extending inward from the peridium; spores brown, pedicellate or with sterigmatal scar" (Zeller, 1949: 52). Sedccula Zeller Tliere is "one genus, Scdecula" (Zeller, 1949: 52) in the family.

Page  105 SCLERODERMA TALES 105 PISOLITI-ACEAE "Fructifications mostly stalked, rarely sessile (in Pisolithus); stems root-like or hard and wood-like; sporocarps subglobose, pear-shaped, or hemispherical; peridium thin or layers not separating readily, breaking away irregularly; gleba dark, made up of roundislh or irregular basidia-bearing sectors or peridioles which loosen and break away at maturity; capillitium none; spores colored, sculptured" (Zeller, 1949: 52). KEY TO (ENERA OF PISOLITIACEAE i. Fructification sessile or with a rootlike stalk; peridium thin....... Pisolithus i. Fructification with a hard woodlike stalk; peridium thick and hard.............................................................Dictyocphalos Pisolithus Albertini and Schweinitz Fructification with a pseudostipe as in some species of Scleroderima or with a stipe more compactly organized; wall of spore case very thin and soon fragmenting to expose the peridioles. The genus is distinguished from Sclerodcrma by the thin wall of the spore case and the presence of peridioles which contain masses of powdery spores. Pisolithus tinctorius (Pers.) Coker and Couch (1'1. XXXVII) Fructification typically large, 5-18(25) cm. high, 4-10(15) cm. in diameter, usually with a thick rooting base, rarely sessile; outer wall very thin and soon breaking up exposing the peridioles; interior yellowish olivaceous becoming brownish or darker; peridioles yellowish (pallid at first according to some authors) becoming watery vinaceous and soon darker, about 1-2 mm. thick and up to 4 mlm. or more long, variously shaped and smaller in the base, separated by a gelatinous dark-colored matrix. Spores 8-12 p in diameter, globose, cinnamon color in mass, verrucose with a lhyaline envelope into which the blunt warts project (mounted in both Melzer's reagent and K(H).

Page  106 1o6 PUFFBALLS AND ALLIES IN MICHIGAN Habit, habitat, and distribution.-This species is not known from Michigan, but should occur here. Discussion.-I have included this most unattractive of all species of fungus in the hope of getting more information on its distribution. It is common in Washington and Oregon along road cuts, where the massive fruiting bodies stick out like halfrotted stumps of large saplings. When fresh it stains everything with which it comes in contact, and when dried the spore powder covers everything in the near vicinity so that it is a disagreeable fungus to handle in any condition. The specimen photographed wIas found in the Olympic National Park, Washington, and represents Pisolithus crassipes, if that fungus is actually distinct from P. tinctorius. Cunningham put the former name in synonymny, and I have followed him here. P. tinctorius has been reported for the Great Lakes region but appears to be rare there. GLISCHRODERMATACEAE "Fructifications subglobose, on a superficial mycelium; peridium simple, thin but hard and tough, opening by an apical pore; gleba with evenly distributed basidia (without sterile veins), capillitium arising from the inner side of the peridium (as in Calvatia); spores sculptured" (Zeller, 1949: 52). Glischroderma Zeller There is "one genus, Glischroderma, found in Europe only" (Zeller, 1949: 52) in the family. TU LOSTOMATACEAE "Fructifications at first hypogeous, sporocarp elevated by the prolongation of a basal tissue into a stout, fibrous, stemlike or cushion-like process; peridium duplex, outer layer partly evanescent, partly remaining as a cuplike volva at the base of the stem, inner layer tlin, dehiscing by an apical pore, several pores, or circumscissilely; gleba without chambers or chambered by the labyrintline separation of the tissues from one another; basidia regularly and evenly distributed in the glebal tissue or forming a rudimentary hymenium on the walls of chambers; capillitium

Page  107 SCLERODERMA TALES 107 well-developed, attached to the inside of the peridium; spores variously sculptured" (Zeller, 1949: 53). KEY TO GENERA OF TULOSTOMATACEAE i. Endoperidium dehiscing circumscissilely or through numerous pores; basidia borne in an elementary hymenium.....................Battarraca 1. Typically not as above............................................... 2 2. Stem more or less readily breaking from spore sac or out of a socket.... 3 2. N ot as above........................................................ 4 3. Spore case opening by an apical stoma...........................Tulostoma 3. Spore case opening by a roughly stellate stoma.................Scliizostoma 3. Spore case opening irregularly.................................... Queletia 4. Exoperidium continuous with outer layer of stem; stipe not volvate...............................................................Phellorinia 4. Exoperidium not continuous with stem; stem volvatc.........Chlamydopus Tulostoma Persoon Fructification at maturity consisting of a spore case, a stalk and a mycelial bulb at base; spore case globose to onion-shaped, with a thin outer coating which is usually deciduous over upper half or adheres as small particles but is more persistent around the base as a sand collar (mycelium binding a rather thick layer of sand), rarely membranous; inner layer smooth or nearly so, membranous, opening by an apical stoma which may be indefinite, definite, fibrillose, tubular, umbonate, or plane; stalk inserted in a socket at base of spore case, woody, smooth to scaly, interior stuffed; gleba of spores and capillitium; capillitium threads attached to wall of spore case, long, septate, branched, and usually the walls thick and almost hyaline in KOH under the microscope; spores variable in shape, smooth or roughened. This is an interesting genus, but apparently examples of it have been rarely collected in Michigan. Potter has collected several species in the region around Ithaca, Gratiot County. KEY TO MICHIGAN SPECIES OF Tulostoma i. Spores with conspicuous more or less parallel ridges; outer layer of spore-case wall membranous and often surrounding the base of the spore sac in the manner of an acorn cup.....................T. striatum 1. Not with above combination of characters............................ 2 2. Stipe more or less covered with coarse, hairlike fibrils.........T. fibrillosum 2. Stipe not with hairlike fibrils......................................... 3

Page  108 1o8 PUFFBALLS AND ALLIES IN MICHIGAN 3. Spore case membrrane smooth..................................T. bruinale 3. Spore case with persistent sand particles held to surface by remains of outer layer....................................................... 4 4. Mouth with smooth edges, round and slightly elevated...........T. sinulans 4. N ot as above........................................................ 5 5. Spores verrucose (under high dry optical system)............... T. campestre 5. Spores appearing smooth under high dry system.............. T. volvulatum Tulostoma striatum Cunningham (P1. XXXVIII, Fig. 1) Fructification 1.5-2.5 cm. high, consisting of a mycelial bulb at the base which contains much sand, a short stipe about 1 cm. long, and an acorn-shaped spore case about 1 cm. broad; spore case at first nearly enveloped by a membranous, pallid, outer wall (exoperidium) which flakes off from upper part but leaves a loose fitting lobed or irregular-margined cup over lower half, the under side of the cup thick because of the amount of agglutinated sand; wall of spore case pallid tan, minutely furfuraceous under a lens; mouth area raised, distinct, fibrillose, opening by a pore; gleba ferruginous salmon; stipe 1-1.5 cm. long, 2-3 mm. thick (when dried), tan to sordid tan (in 1 specimen it is paler than the spore case), longitudinally striate. Spores 6-8x4-7 p in diameter, ellipsoid, ovoid, angular or subglobose, rusty yellow in iodine, pallid yellowish in KOH, marked by parallel or nearly parallel ridges extending almost around the spore, and then doubling back by a U-shaped bend, ridges in some specimens convergent toward the poles; capillitium of tlireads 4-8 p in diameter, slightly enlarged at the septa, walls thickened (very irregularly so in main branches), liyaline in KOH, pale yellowish in iodine. Habit, habitat, and distribution.-Until 1948 the only North American collection of this species known to me was one made October 24, 1943, by F. K. Sparrow in sandy soil on the H. V. S. Ogden tract, near Dixboro, WXashtenaw County. During the season of 1948, mostly in November, Potter collected it in quantity in abandoned fields near Riverdale, Gratiot County. To judge from his collections it was second in abundance there only to T. campcstre.

Page  109 SCLEROD ERMA TALES log Discussion.-Thel c(onspicuously ridged spores clearly distinguish this fungus from T. poculatum, in which it would otherwise be placed. Since this is the main difference between the 2 species, a microscopic examination is necessary to distinguish them. Potter took the following notes on fresh specimens of striatum: "Spore case when fresh and moist 'dark indian red' or red brown (338-3), occasionally mineral brown (339-3), fading rapidly to pallid tannish and becoming minutely furfuraceous; mouth area pale tan from first and sharply delimited in color. Stipe 2-5 mm. thick, pure white, becoming pallid or finally pallid tan in age, smooth." Tulostoma fibrillosum VWhite (P1. XXXVIII, Fig. 2) Fructification consisting of a bulbous base 6-8 mm. in diameter, a stipe 5-8 cm. long and 3-5 mm. thick when dried, and a spore case 10-15 mm. broad; spore case globose to depressedglobose, whitish to ashy, smooth to obscurely pitted, sand usually adhering on under side as a basal zone, original outer covering of spore case sometimes remaining as a few small brownish flecks; mouth area indistinct, opening by a pore, with a lobed to fibrillose margin; stipe pulling away from sand case to leave a narrow circular free zone between it and spore case; buried part of stipe at first covered by a loose copious to sparse fibrillosity with rather numerous coarser fibrils extending some distance into sand; surface with a dull ochraceous tawny cortex which bleaches to white or ashy when exposed and soon scales off after first becoming areolate, pallid beneath. Spores ellipsoid, ovoid to globose or more irregular, 5-7 or 5-7 x4.5 p with a thin hyaline envelope into which minute warts project; capillitium of hyaline threads with thick hyaline walls in KOH, yellowish in iodine, swollen at septa, 4-7 p in diameter. Habit, habitat, and distribution.-Gregarious to scattered in sand. The description is based on a collection made by Morten Lange at Sleeping Bear Dune, Leelanau County, July 24, 1947. It is the only typical material I have seen from Michigan.

Page  110 110 PUFFBALLS AND ALLIES IN MICHIGAN Discussion.-The comment by Lloyd that this name applies to a condition rather than a species may be true, but I illustrate the species here in order that it may be easily recognized if found. In the characters of the mouth and the spore case, and in such microscopic features as the spores and capillitium, it is obviously very close to T. campestre. One point should be kept in mind, however, by those who ascribe the differences to habitat. Both T. campestre and T. fibrillosum occur in sand and the former has a stipe practically free of fibrils, whereas the latter is distinctly fibrillose. This character may become obscured in herbaria, for the stems may finally become more or less polished from handling. Long (1946: 175-76) has given a detailed account of the type. Tulostoma campestre Morgan (1'1. XXXIX, Fig. 1) Fructification stipitate with a depressed-globose head; head (spore case) 10-20 mm. broad, 10-15 mm. high, outer layer of wall containing much sand and breaking up into small, areolate to almost granular, subpersistent scales consisting of sand particles held to surface by remains of outer layer, these scales finally deciduous at least over the upper part revealing the smooth inner surface, region around stipe attachment (or whole underside) covered by a persistent sandy coating not separating from spore-case wall as a membrane; spore-case wall pale tan to ashy gray depending on degree of weathering; mouth area a somewhat distinct spongy, somewhat elevated disc and opening by a pore, the margins of which vary from lobed to lacerate; stipe 2-5(6) cm. long, 2.5-4(5) mm. thick when dried, usually with a marginate basal mycelioid bulb; surface more or less pale rusty brown, longitudinally striate when dried, the brown covering often broken up into lacerate scales and shrinking away from spore case in drying so that the latter extends down and around point of attachment as a collar; interior whitish. Spores 4.5-6.2 p in diameter, globose to subglobose, quite irregular in shape; verrucose with a layer of hyaline material collapsed around the warts; near ochraceous tawny in iodine;

Page  111 SCLERODERMA TALES 111 capillitium of thick-walled threads up to 7 p in diameter, slightly enlarged at the septa, hyaline in KOH, only faintly yellowish in iodine. Habit, habitat, and distribution.-Single to gregarious in sandy fields. Kauffman reported the species as probably occurring throughout the state, but Potter appears to have been the first person to make extensive collections of it. It fruits during the summer and fall, but as the fructifications are rather hard after drying they persist a long time. Discussion.-The description is based upon material identified by Lloyd. The manner in which the sand adheres to the spore case is a distinctive feature of the plant. Tulostoma brumale Persoon Fructification subglobose, 1-1.5 cm. high, 1-2 cm. in diameter, the brown outer peridium soon retreating, leaving the inner peridium smooth and membraneous; mouth short-tubular, entire, prominent; collar inconspicuous; stipe usually pallid, slender, lacerate-scaly or nearly smooth, with a small mycelial bulb stuffed with loose silky threads, 1-5 cm. long, 2.5 mm. thick, capillitium 4-7 p thick, septate somewhat swollen at the joints, light-colored, hyaline; spores subglobose, 3-5 P in diameter, minutely verrucose, some with short pedicels. Habit, habitat, and distribution.-In sandy soil. Longyear reported the species as not common in the state. Discussion.-I have seen no material of this species. The description has been adapted from White (1901: 429). Longyear reported the fungus under the name Tylostoma mammosum Fr. White used the name Tulostoma pedunculatum, but according to the International Rules the correct name is T. brumale. Tulostoma sinmulans Lloyd (P1. XXXIX, Figs. 2-3) Fructification 2-4 cm. high, consisting of a bulbous to irregular mycelial basal mass of fibrils and sand, a stipe 1.5-3 cm. long and 3-4 mm. thick, and an acorn-shaped to globose spore case 10-15 mm. across the base; spore case dark reddish brown

Page  112 112 PUFFBALLS AND ALLIES IN MICHIGAN when fresli, b)tt so covereld witl sand and cartl as to obscure all details, witll a thin weft of fibrils at first covering tle debriscoated surface, debris slowly wearing away to expose the brownish to pallid spore-case wall, mouth small, round, edges smooth, slightly and abruptly elevated, area surrounding the mouth at first concolorous with remainder of wall, but in extreme age becoming grayish and hence darker; stipe with a surface covering of rusty brown, appressed to lacerate scales, paler but dull brownish between, becoming grayer to pallid in age; inside pallid; some longitudinally striate wzhen dried, spore case slhrinking away from stipe in drying leaving a collar. Spores 4-6 p in diameter, globose to subglobose, surrounded by a gelatinous layer into which plugs of material project to produce a finely tuberculate appearance; sand color in mass, under tlhe microscope pale yellowish; capillitium of flexuous hyaline tlreads witlh thick walls, 4-6 p in diameter, occasionally branched, a few cross walls present, and joints in some of the tlhreads slightly swollen. Habit, 1habitat, and distribution.-GreCgar-ious on sandy soil at edge of white pine groves near Ithaca, Gratiot County, and Carson City, MIontcalm County. Potter collected this species in some quantity. )iscussioti.-Potter found overwintered material in July and by visiting the same localities later finally obtained excellent fresl material from November 1 to 15. The only fruiting bodies whlich slowed any tendency to darken around the mouth were tlose which overwintered in the field. Tulastoma volvulatun sensu Coker and Couch "Peridiun up to 1 cm. thick, covered until age with a rather thin, brown, sandy coat wllicll wears away slowly and about e(qually over the surface, exposing the nearly white inner peridium. Moutl plane, not regularly circular, margin clean and smootlh, single in all nine plants of our collection. There is no cortical cup at the base of the peridium. Collar short, fimbriate. Stem 2-3 cm. long, 1.5-3 mm. thick, witlh a brown, more or less scaly b)tt not conspicuously lacerate surface, cespitose at times, in which cases tlhe peridia may be fused.

Page  113 SCLERODERMA TALES 113 "Spores subsplerical, irregular, somewhat angled, practically smooth (even under oil immersion), 4.2-5.5 x 5-7.4 p. Capillitium threads very irregular, nearly hyaline, paler than the spores, up to 5 P thick, with walls of varying thickness" (Coker and Couch, 1928: 155). Habit, habitat, and distribution.-Gregarious on sand in an abandoned field, Riverdale, Gratiot County. Disclussioln.-One collection by Potter (Potter-6776) appears to belong here. It consists of 4 fruiting bodies. The spore case of tie largest is 7 mm. broad and the stipe lox 1.5 mm. The stipe is wlhitish beneath a thin coating of a wood-brown color which tends to scale off. The spores are as Coker and Couch described them, that is, irregular, smooth under ordinary magnifications, and about 5-6.5 p in diameter. Coker and Couch regarded their material as representing a slender form of the species. I have not been able to examine authentic specimens of the species, and consider the identification of the Potter collection tentative. ASTRAEACEAE "Fructifications epigeous or at first lhypogeous, sessile; peridium of several layers, the outer two or three becoming the very heavy exoperidium, which dehisces stellately; the endoperidium thin, membranous; columella none; gleba separated by delicate sterile veins into basidia-bearing sectors in which the basidia are regularly distributed throughout the context; basidia broadly club-shaped; spores spherical, sculptured; capillitium none.2 "One genus, Astracus" (Zeller, 1949: 53). Astraeus Miorgan "Mycelium fibrous, proceeding from all parts of the surface. Peridium subglobose, composed of two persistent coats; outer peridium thick, coriaceous-cartilaginous, at first concrete with the inner peridium, then at maturity burst into segments and torn away; inner peridiuin tlin, ecmbranaceous, sessile, de2 But see generic description and that of A. Iiygromictricus.

Page  114 114 PUFFBALLS AND ALLIES IN MICHIGAN hiscent at the apex by a single mouth. Columella none. Capillitium originating from the inner surface of the pcridium; threads long, much branched and interwoven; spores large, globose, minutely warted, brown." The original description of the genus by Morgan is quoted. Astraeus differs from Gcastrum, from which it was segregated by Morgan, in the lack of open chambers in the gleba, and consequently no organized hymenium, and by the long, muchbranched capillitial tlreads which are scarcely different from the elemental hlyphae of the peridium and continuous witl it. Tlhe columella is lacking and the spores much larger than in Geastrunm. Tlhe genus is represented in Michigan by 1 species. Astracus hygrometricus (Pers.) Morgan (PI. XL, Fig. i) Fructification 1-2(3) cm. in diameter, globose to depressedglobose wlien young; arising from black hairlike rhizomorphs some of which may adllere to the base as appressed hairs; outer wall tliick, splitting into 7 to 15 pointed rays which expand (bend backward so as to expose spore sac) when wet and curve in again over spore sac whecn dry, that is, they are hygroscopic; the outer waall distinctly layered, outer layer (under surface of expanded rays) composed of a thin matted-fibrillose layer with sand adhering to it or embedded in it, interior to this is a fibrous layer wlhiclh is quite hard in dried plants, the next layer is that on the exposed upper surface of expanded rays and is deep purple drab, thick, and variously rimose; between the rays and tlhe spore sac in freshly matured specimens is a thin pallid papery layer usually visible as fragments around base of the spore sac; spore sac sessile, globose to depressed-globose, 1-2 cm. broad, wall pliable, matted-fibrillose or somewhat fibrillosereticulate, usually pallid grayish when collected; opening by a slit or tear or with an irregular pore (no characteristically formed mouth); gleba wlhite when young, cocoa brown at maturity. Spores 7-10.5 p, globose, with a very thin hyaline envelope; interior to tllis envelope is a tlickened colored wall with many

Page  115 SCLERODERMA TALES 115 pores (both round and irregular) so that the outer surface of the colored wall does not appear entirely smooth; hyaline plugs extending through pores into hyaline envelope and at maturity the plugs persist as hyaline or yellowish warts; capillitium as in most species of Geastrum, that is, of pallid to yellowish thickwalled cells with incrusting material, but is as a rule more branched. Habit, habitat, and distribution.-In colonies, gregarious to scattered, developing near the soil surface and becoming exposed at maturity. It is a characteristic species of waste sandy areas throughout the state. Old fruiting bodies may be found any time during the year, but freshly matured specimens are to be found during the fall months only. It is then one of the commonest of Michigan puffballs. Large numbers can frequently be collected in abandoned sandy fields, as those in the Waterloo Area, Jackson County, in the lake region in northwestern Washtenaw County, and in similar areas. Discussion.-The manner in which the fruiting bodies open and close as they become wet or dry is an interesting feature and one which has always attracted much attention. This, in addition to the habitat, matted-fibrillose surface of the spore sac, rimose inner (or upper) surface of the rays, and cocoacolored gleba are the field characters to learn. The structure of the spore is very interesting though details cannot be made out with ordinary optical equipment. The description of the spores given is based upon an examination made with an oil-immersion apochromatic objective (2 mm. N. A. 1.3). The interesting feature is that the highly colored, thick spore wall is punctured with numerous pits which are merely large enough so that the outer surface of the colored wall appears slightly uneven, not warty. The warts are produced by the drying down or solidifying of the hyaline material extruded through the pores of the colored wall. The hyaline material, in immature spores, forms an even envelope over the colored wall; hence immature spores are smooth or nearly so, and very old spores roughly treated may have most of the warts knocked off. In order to evaluate properly the spore markings as taxonomic

Page  116 116 PUFFBALLS AND ALLIES IN MICHIGAN characters in the Gasteromycetes, one must study carefully the origin of such markings, for it is likely to shed light on the manner in which the markings vary. CALOSTOMATACEAE "Fructifications epigeous or at first hypogeous, stalked with a root-like, lacunose basal process; peridium duplex, exoperidium cartilaginous, extended below into a rootlike stalk and often cupulate around the base of the sporocarp; endoperidium cartilaginous, with an ornate apical stellate stoma below which the spore sac is suspended; gleba pulverulent; spores spherical or ellipsoid, smooth or sculptured" (Zeller, 1949: 53-54). Calostoma Zeller There is "one genus, Calostoma" (Zeller, 1949: 53-54), in the family. NIDULARIALES "Fructifications small, sessile, cupulate, campanulate or depressed globose; peridium of one to four layers, dehiscing by rupture of an epiphragm or lid over the top, or when this is absent, by irregular fissuring of the wall; gleba enclosed in one or many globose or lens-shaped peridioles; peridioles attached to the inner wall of the peridium by a mucilaginous secretion or by threadlike funiculi, escaping singly or they may be forcibly ejected from the exoperidium; capillitium none" (Zeller, 1949: 54)These fungi are popularly known as the "bird's-nest fungi," because the fructification with the peridioles reminds one of a bird's nest with eggs in it. It is hard for some people to accept the idea that these plants can be even remotely related to such fungi as Calvatia gigantea. KEY TO FAMILIES OF NIDULARIALES i. Exoperidium more or less urceolate and firm at maturity; glebal chambers (peridioles) egglike, remaining attached or free within outer peridium, empty......................................... Nidulariaceae i. Exoperidium collapsed at maturity; single, spherical, glebal chamber violently discharged at maturity, filled with gel or gelatinous tissue......................................................... Sphaerobolaceae

Page  117 NIDULARIALES 117 NIDULARIACEAE "Fructifications epigeous, with hard peridium which opens cuplike at maturity; gleba with a few mostly flattish, rounded, closed chambers (peridioles), the hard walls of which are lined with basidial hymenium and in the mature fructification are isolated or freed from the cup-like fructification by ejection or by the deliquescence of the intervening tissue" (Zeller, 1949: 54). KEY TO GENERA OF NIDULARIACEAE i. At least some peridioles attached to fructification by a cord (funiculus).. 2 1. Peridioles not attached, instead embedded in mucilage............... 3 2. Wall of fructification thick but consisting of a single layer; peridioles whitish from a thick tunica.................................. Crucibulum 2. Wall of fructification 3-layered; tunica of peridioles thin, gray to black............................................................... Cyathus 3. Fructification without an epiphragm; the wall thin and fragmenting at m aturity....................................................N idularia 3. Epiphragm present; fructification persistent.......................Nidula Cyathus Persoon Fructification shaped like a vase or an inverted bell, point of attachment truncate and narrow, wall of 3 layers, apex at first closed by a thin membrane (epiphragm); peridioles lenticular, attached to wall by a funiculus; spores hyaline, smooth, usually thick-walled. The 3-layered wall, which is one of the main characters of the genus, is not always clearly evident. The epiphragm and persistent funiculi are also important. Michigan has 3 species. Cunningham recognized 5 in Australia. KEY TO MICHIGAN SPECIES OF Cyathus i. Inner surface of fructification striate at least near upper edge......C. striatus 1. N ot as above........................................................ 2 2. Spores 20-35, long..........................................C. stercoreus 2. Spores 8-12(15)IU long............................................. C. olla Cyathus striatus Persoon Fructification 10-15 mm. high, 2-3 mm. broad at point of attachment, 5-10 mm. broad at apex, attached to substratum

Page  118 118 PUFFBALLS AND ALLIES IN MICHIGAN by a pad of cinnamon-brown mycelium, at maturity vase-shaped to somewhat trumpet-shaped, exterior coarsely fibrillose or in unopened vases almost fibrillose scaly, color pale to dark cinnamon brown or sometimes grayish brown (apparently from weathering), sometimes slightly striate above when hairs are more or less worn off; mouth at first closed by a thin epiphragm which is at first covered over by a shaggy-fibrillose covering, breaking and disappearing at maturity; inner surface glabrous, shining, distinctly striate, pallid to nearly black but usually some shade of drab or purplish drab; peridioles about 1-2 mm. in diameter, flattened (more or less disc-shaped or angular from pressure), usually occupying lower half of cup, usually attached to cup by an elastic cord; spores (12)15-20(22) x (6)8-12 p, hyaline, thick-walled, but wall becoming thinner as they mature. Habit, habitat, and distribution.-Subcespitose to gregarious on sticks, bark, and ligneous and vegetable debris generally. It is a much more common species in the state than the number of collections preserved in herbaria would indicate. Discussion.-This is a very interesting little fungus easily recognized in the field by the brown, shaggy-fibrillose covering of the exterior and the conspicuously striate, glabrous interior. In drying, however, the striations often become obscured so that they are not conspicuous in herbarium specimens. Cyathus stercoreus (Schw.) de Toni Fructification 5-10(15) mm. high, 4-8 mm. broad at top, 1-3 mm. at point of attachment and with or without a basal brownish mycelioid pad, vaselike to goblet-like in shape, outer surface densely fibrillose, color of fibrils variable, whitish, cinnamon buff, or darker brownish, sometimes brownish around base and whitish over upper expanded part, fibrils gradually weathering away so that in age the outer surface may be nearly smooth and in some with depressed circular zones; inner surface smooth, shiny, pale to dark lead color; epiphragm present but soon breaking and all traces vanishing; peridioles 1-2 mm. in diameter, black, compressed, attached to cup by a funiculus (those on top not always attached), smooth and shiny; spores (22)25-30

Page  119 NIDULARIALES 119 (35) x (18)20-27(35) i, nearly hyaline, subglobose to nearly ellipsoid, thick-walled. Habit, habitat and distribution.-Cespitose to scattered on dung, manured ground, or around edges of sawdust piles. Rather common in Michigan in early summer or in the fall. Potter (Potter-4254) found it on old sawdust in a meadow near Ithaca, Gratiot County. Such a habitat is generally regarded as unusual. Discussion.-The fibrils extend up to and beyond the edge of the fruiting body to create a fimbriate appearance at first, but as the fibrils disappear the edge becomes even and glabrous. The lack of striations in the inner wall of the cup serve to distinguish this species from C. striatus. Cyathus olla Persoon Fructification 8-12 mm. high, 5-10 mm. wide at top, substipitate and about 2 mm. broad at base (2-7 mm., according to White), in age flaring to recurved at top, when young with margin incurved and goblet-shaped to inverted bell-shaped, outer surface brown (cinnamon brown to paler and about cinnamon buff), shaggy fibrillose young, merely appressed fibrillose to nearly glabrous at maturity and then obscurely zoned, fibrils yellowish brown, inner surface smooth, silvery brown to lead color, peridioles large, disc-shaped, 2-3 mm. wide and o.5 mm. thick in center, dull brownish, attached to cup by a strong whitish thread (funiculus); spores 8-12(15) X(6)8-12 p, thickwalled, hyaline, ovoid to ellipsoid. Habit, habitat, and distribution.-The only material from this state which I have seen was collected by M. Weideman on a piece of wood in a greenhouse in Ann Arbor, Washtenaw County, May 26, 1893. Discussion.-The small spores distinguish it from the other species included here. It is most likely to be confused with C. stercoreus, but at maturity it has a much more flaring margin, and the peridioles are not black.

Page  120 120 PUFFBALLS AND ALLIES IN MICHIGAN Crucibulum Tulasne Fructification short cylindric to cyathiform, sessile, wall one layer thick, exterior fibrillose to velvety, apex covered by an epiphragm; peridioles pale-colored, attached by a funiculus; spores hyaline, smooth. Michigan has only i species, the description of which follows. Crucibulumn levis (D.C.) Kambly and Lee (P1. XL, Fig. 2; P1. XLI) Fructification 5-10(12) mm. high, 5-o1 mm. across the top and 3-5(7) mm. wide at base, when young subglobose, cupshaped, or short-cylindric, at maturity narrowed slightly from the top toward the sessile truncate base which is attached to a subiculum; surface velvety to appressed silky but almost smooth in age; tawny yellow becoming nearly cinnamon brown, in age somewhat sordid; inner surface smooth, shiny, whitish to gray (pallid cinnamon, according to Cunningham); margin erect or slightly flared, even, thick, entire, sometimes slightly contracted; wall of a single layer of woven hyphae; epiphragm soon disappearing; peridioles pallid (pallid brown according to Cunningham), lenticular, circular in outline, 1-2 mm. broad, attached to cup by a cordlike strand (funiculus); spores (4)7-10 x4-6 p, hyaline, thick-walled (wall about 1 p, according to Cunningham). Habit, habitat, and distribution.-Gregarious, scattered, or solitary on ligneous or vegetable debris. A common species during favorable weather in the summer and fall. It has been collected mostly in the southern part of the state, but this should not be regarded as indicative of its distribution. Discussion. —This species is known the world over under the name Crucibulum. vulgare Tulasne, but since species names cannot be conserved according to the International Rules, it is necessary to use the combination given above. Cunningham stated that there is no evidence that the Cyathus levis De Candolle was the same as C. vulgare. This is apparently an instance in which interpretation is difficult, for others believe the oppo

Page  121 NIDULARIALES 121 site. As the epithet levis must be accounted for, the solution adopted here seems to me to be the most sensible one. SPHAEROBOLACEAE "Fructifications tiny, spherical at first; peridium of several layers, of which the second inner layer is formed of turgescent cells appearing as a radial palisade; gleba of basidia-bearing sectors separated by sterile veins, or of basidia-bearing cavities formed by the splitting of tissues; the gleba becomes gelatinized at maturity and is ejected as a whole from the peridium" (Zeller, 1949: 55). KEY TO GENERA OF SPHAEROBOLACEAE i. Basidia borne irregularly throughout the basidia-bearing sectors.................................................................. ph aerobolus i. Basidia borne in hymenia on the walls of cavities............. idulariopsis Spliaerobolus Persoon Fructification small, subglobose, 4-layered, dehiscing by stellate rupture of exoperidiunm, evagination of the endoperidium and forcible discharge of the single viscid glebal ball; spores smooth and hyaline. Only 1 species is recognized from Michigan, but S. iowensis should also occur in the state. Sphaerobolus stellatus Persoon (P1. XLII) Fructification small, 1.5-2.5 mm. in diameter, developing in the substratum but so near the surface that its upper surface soon breaks through and is exposed, dull ochraceous to pallid at this stage, soon opening by a number (5 to 9) of lobes giving appearance of a small cup; thin receptaculum now suddenly reversing itself outward, throwing the flattish glebal ball several feet, when reversed the receptaculum appears as a watery-white sphere situated on the lobes of the outer wall; glebal ball slippery, smooth, very dark chestnut brown in age; spores 7.5-10 x 3.5-5 P.

Page  122 122 PUFFBALLS AND ALLIES IN MICHIGAN Habit, habitat, and distribution.-Densely gregarious on rotten wood, packed sawdust, and old dung of horses. It is fairly common in the southeastern part of Michigan on the packed sawdust around old sawdust piles. It fruits most abundantly early in June. Discussion.-In many respects this little fungus is truly a small cannon. Miss Walker calculated that it can throw its glebal ball to a height of 14 feet. Walker and Anderson discovered that the force for the discharge was obtained from the difference in osmotic pressure of the glycogen (originally present) and the sugars into which the glycogen was suddenly changed. The glebal ball contains cells rich in food stuffs in addition to the spores and the entire ball can germinate, that is, hyphae develop from the cells just mentioned (called gemmae) as well as from the spores. The fungus is not a spectacular sight in the field, but a biology teacher can have much fun with it in the laboratory. PODAXALES "Fructifications epigeous, stalked or with percurrent columella, pileate at maturity, angiocarpic; stipe long or short, continued to the apex of the fructification as a columella; peridium simple or 2-3-layered at maturity, left at maturity in part as pileus, or volva, or annulus on the stem; gleba at first with hymenium of basidia covering the walls of chambers or pores or lamellae, persistent or pulverulent; capillitium wanting (except in Podaxis); spores colored" (Zeller, 1949: 55). KEY TO FAMILIES OF PODAXALES i. Gleba not powdery at maturity.............................. SECOTIACEAE 3 i. Gleba powdery at maturity...................................PODAXACEA SECOTIACEAE "Fructifications mostly epigeous, stalked or sessile, at first companulate, like an agaric button, angiocarpic; stalk continued above as a percurrent columella; peridium mostly as a cap 3 The Secotiaceae is the only family of this order in Michigan, and its members will be found to have a powdery gleba at maturity.

Page  123 PODAXALES 123 covering the gleba, free at maturity or opening by a transverse slit; gleba chambered or with irregular pores or with anastomosing, lamelloid, tramal structures, dark brown to blackish, sometimes with cystidia, not becoming a powdery spore mass at maturity; spores dark, smooth or sculptured; capillitium none" (Zeller, 1949: 55-56). KEY TO GENERA OF SECOTIACEAE i. Stems not volvate, not annulate; gleba brown.......................... 2 i. Stems volvate and/or annulate; spores black to dark brown............. 3 2. Fructifications short-stemmed or sessile; pileus mostly ovoid, obtusely conic to globose or depressed-globose...........................Secotium 2. Fructifications with long slender stems; pileus cylindric to conic.. Galeropsis 3. Gleba lamelloid; stem volvate; gleba free from expanded stipe apex...............................................................M on tagnea 3. Gleba of anastomosed lamellae and attached to expanded upper part of stem (the pileus); stem annulate................................Longula 3. Volva and annulus both typically present; gleba anastomosed to poroidlamellate.............................................. Gyrophragmiurn Secotium Kunze Fructification stipitate; spore case with wall one layer thick, its surface smooth, fibrillose or viscid, sometimes scaly in age, variously colored, margin at first intergrown with stipe tissue but in some species soon separating; stipe central, extending through the gleba as a columella; gleba of persistent tramal plates attached to spore-case wall or upper part of columella (reminding one of lamellae), but anastomosed to enclose vesiculose to elongated cavities; spores colored, smooth, typically with an apical pore. This is a relatively large genus. Cunningham reported fourteen species from Australia, and it should be represented in Michigan by more than the 2 species which follow. S. pingui, a common fungus of the pine forests of the Cascade Mountains, should occur in the jack and red pine forests in northern Michigan. Secotium agaricoides (Czern.) Holl6s (P1. XLIII) Fructification 1-6 cm. wide, 1.5-7 cm. high, consisting of a short stem and a subglobose, conic, heart-shaped or convex

Page  124 124 PUFFBALLS AND ALLIES IN MICHIGAN umbonate spore sac, margin of spore sac confluent with upper part of stem, stem extending as a reduced structure (the columella) to apex of spore sac; surface of fructification pure white when young and fresh, glabrous to unpolished, in age discoloring slightly to yellowish or pale leather color and in some specimens becoming more or less fibrillose-scaly from the separation of fibrils into fascicles, in age tending to shred or tear especially near the zone of attachment to the stipe; gleba fleshy and white when young, becoming yellow and finally yellowish brown or darker, coarsely cellular to labyrinthiform, the tramal plates forming lamellae and sparsely anastomosed, vertically arranged and attached to the spore-sac wall. Spores 7.8-9.3 x 5-7, smooth, oval to elliptic, with a thin hyaline envelope, a thick yellow inner wall possessing an apical pore, and a basal stump of a pedicel. Habit, habitat, and distribution.-Cespitose, gregarious, or scattered usually in pastures and cultivated fields. It fruits during the summer and fall. Johnson listed it as fruiting from Maya to October. Excellent specimens from Michigan were collected in Washtenaw County as early as 1893. The collector's name was not given, nor date other than the year. C. D. LaRue found some fine specimens on a rotten log on the south side of Monroe Lake, Cheboygan County, on July 29, 1929; and Victor Potter found luxuriant fruitings of numerous fruiting bodies in clusters in September near Ithaca, Gratiot County. Discussion.-One's first impression of this fungus is that it is a mushroom which has failed to expand. In the fleshy, white, immature stage it is edible. The fungus is apparently fairly rare in Michigan. Because of its unusual features, however, collectors nearly always bring it in; hence, the collections in herbaria are likely to be more numerous for it than for some other more common but less interesting species. Secotium coprinoides Routien Fructification 4 mm. high, white, consisting of a stalk and a pileus-like upper part nearly 2 mm. in diameter, the upper fertile region united to the stalk only near the apex of the latter

Page  125 LITERATURE CITED 125 and consisting of about 18 glebal chambers; peridium white, of filamentous-inflated cells 25-35 p thick. Hymenium at maturity consisting of basidia and paraphyses; basidia 2 to 4 spores; spores ellipsoid, smooth, black, 18(23.5)-30.5 x 12.6(12.75)-16 p, pedicellate, pedicel 2x 1.5 P, 2, 3, or 4 spores not uncommonly grown together and united into a group. Habit, habitat, and distribution.-Specimens of this fungus developed on soil brought into the laboratory from woods near East Lansing, Ingham County. Discussion.-The descriptive data are taken from Routien (1940). It would be very interesting to learn more about this minute species and make a careful study of its spores in relation to those of some of the species of the western United States. Routien did not illustrate a germ pore, apical or subapical, in his study, and since none was mentioned in his description I assume that this is one character in which the spores of Coprini differ. PODAXACEAE "Fructifications angiocarpic, like agaric buttons, epigeous at maturity, clavate, ovoid or fusiform, stalked or almost sessile; stem firmly fibrous, extending percurrently as a columella; peridium pileate, simple or plicately scaly, brittle, easily splitting margin loosening from the stipe at maturity or opening by longitudinal splitting; gleba at first chambered by anastomosing tramal tissues or lamelloid tramal plates, powdery at maturity; basidia persisting in Podaxis; capillitium well developed as elaters, or wanting; spores dark, smooth" (Zeller, 1949: 56). KEY TO GENERA OF PODAXACEAE i. Fructification sessile or nearly so; gleba with capillitium......Endoptychum i. Fructification with long stipe; gleba with capillitium (elaters)........Podaxis LITERATURE CITED BULLER, A. H. R. 1922. Researches on Fungi. 2:i-xii+ 1-492. COKER, NV. C., and J. N. COUCH. 1928. The Gasteromycetes of the Eastern United States and Canada. Chapel Hill, N. C.: Univ. North Carolina Press. Pp. i-ix+ 1-201, 123 pls.

Page  126 126 PUFFBALLS AND ALLIES IN MICHIGAN CUNNINGHAM, G. H. 1942. The Gasteromycetes of Australia and New Zealand. Dunedin, N. Z.: Privately published. Pp. i-xv, 1-236, pls. i-xxxvii. DODGE, C. W. 1931. Alpova, a New Genus of Rhizopogonaceae, with Further Notes on Leucogaster and Arcangeliella. Ann. Mo. Bot. Gard., 18: 457-64. DODGE, C. V., and S. M. ZELLER. 1934. Hymenogaster and Related Genera. Ann. Mo. Bot. Gard., 21: 625-708. FITZPATRICK, H. M. 1913. A Comparative Study of the Development of the Fruit Body in Phallogaster, Hysterangium, and Gautieria. Ann. Mycol., 11: 119-49. HOLL6S, L. 1904. Gasteromycetes Hungariae. Die Gasteromyceten Ungarns. Leipzig. Pp. 1-278, 31 pls. JOHNSON, MINNIE MAY. 1929. The Gasteromycetae of Ohio: Puffballs, Birds'-Nest Fungi and Stinkhorns. Ohio Biol. Surv. Bull., No. 22., Vol. 4, No. 7: 271-352, 5 pls. KAMBLY, PAUL E., and ROBERT E. LEE. 1936. The Gasteromycetes of Iowa. Univ. Iowa Studies. Studies Nat. Hist. N.S., No. 326, 17, No. 4: 121-85. KAUFFMAN, C. H. 1908. Unreported Fungi for 1907, with an Outline of the Gasteromycetes of the State. Rept. Mich. Acad., o1: 63-84. LLOYD, C. G. 1898-1926. Mycological Writings, Vols. 1-7. Cincinnati, Ohio: Privately published. LONG, W. H. 1946. Studies in the Gasteromycetes. XIII. The Types of Miss White's Species of Tylostoma. Mycologia, 38: 171-79. LONGYEAR, B. 0. 1904. A Preliminary List of the Saprophytic Fleshy Fungi Known to Occur in Michigan. Rept. Mich. Acad., 4: 113-24. MCILVAINE, C., and R. K. MACADAM. 1912. One Thousand American Fungi. New ed.; Indianapolis: Bobbs-Merrill Co. Pp. i-xxvii, 1-749, ccvii pls. ROUTIEN, JOHN B. 1939. Observations on Gasterella lutophila. Mycologia, 31: 416-17. --- 1940. Two New Gasteromycetes. Ibid., 32: 159-69.

Page  127 LITERATURE CITED 127 SHANTZ, H. L., and R. L. PIEMEISEL. 1917. Fungus Fairy Rings in Eastern Colorado and Their Effect on Vegetation. Journ. Agric. Res., 11: 191-245, 30 pls. SWARTZ, DELBERT. 1933. Some Developmental Characters of Species of Lycoperdaceae. Amer. Journ. Bot., 20: 440-65. WHITE, V. S. 1901. The Tylostomaceae of North America. Bull. Torrey Bot. Club, 28: 421-44. ZELLER, S. M. 1939. Developmental Morphology of Alpova. Oregon State Monogr., Studies in Bot., No. 2: 1-19. --- 1947. More Notes on Gasteromycetes. Mycologia, 39: 282-312. -- 1948. Notes on Certain Gasteromycetes, Including Two New Orders. Ibid., 40: 639-68. --- 1949. Keys to the Orders, Families, and Genera of the Gasteromycetes. Ibid., 41: 36-58. ZELLER, S. M., and C. W. DODGE. 1918. Rhizopogon in North America. Ann. Mo. Bot. Gard., 5: 1-36. --- 1929. Hysterangium in North America. Ibid., 16: 83-128. --- 1937a. Elasmomyces, Arcangeliella, and Macowanites. Ibid., 23: 599-638. --- 1937b. Melanogaster. Ibid., 639-55. ZELLER, S. M., and LEVA B. WALKER. 1935. Gasterella, a New Uniloculate Gasteromycete. Mycologia, 27: 573-79.

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Page  129 INDEX TO GENERA AND SPECIES Abstoma 81 Alnus 18 Alpova 3, 15, 17, 126, 127 cinnamomeus 17 Aporophallus 30 Arachnion 37 album 37 Arachniopsis 39 Araneosa 37 Arcangeliella 21, 22, 126, 127 asterosperma 22 Aseroe 30 Astraeus 82, 83, 113, 114 hygrometricus 5, 113, 114 Pteridis 5 Battarraea 10, 97, 107 Blumenavia 30 Boletus parasiticus 1oo Bovista 9, 39, 72, 74, 75, 76, 81 minor 76, 78 pila 6, 76, 77, 78 plumbea 75, 76, 77, 78, 79 Bovistella 9, 38, 62, 72 atrobrunnea 72, 74, 75 echinella 72, 73, 74 radicata 7, 72 Bovistina 81 Bovistoides 39 Broomeia 79 Calbovista 79 Caloderma 98 Calostoma 1 16 Calvarula 29 Calvatia 7, 8, 9, 11, 39, 40, 46, 62, 1o6 Bovista 40, 41, 44, 45 caelata 44, 45 craniformis 11, 40, 43, 44, 45 cyathiformis 11, 39, 40, 41, 44 elata 46 fragilis 40, 41, 42 gigantea 9, 11, 40, 42, 44, 45, 16 saccata 46 var. elata 40, 45, 46 Castoreum 81 Chlamydopus 107 Chondrogaster 15 Clathrogaster 25 Clathrus 30 Claustula 30 Clavaria 64 Colonnaria 30 Colus 30 Corditubera 15 Cortinarius 63 Cremeogaster 15 Crucibulum 117, 120 levis 120 vulgare 120 Cyathus 11 7 levis 120 olla 117, 119 stercoreus 117, II8, 119 striatus 117, 119 Dictyocephalos 105 Dictyophora 30, 34 duplicata 8, 34, 35, 36 indusiata 35 multicolor 35 Diplocystis 79 Disciseda 9, 38, 68, 71 candida 69, 71 Muelleri 69, 71 subterranea 69, 70, 71 Echinophallus 30 Elasmomyces 21, 127 Endoptychum 125 Floccomutinus 30 Galeropsis 123 Gasterella 13, 14, 127 lutophila 14, 126 Gasterellopsis 13, 14 silvicola 14 Gastrosporium 25 Gautieria 23, 126 graveolens 24 morchelliformis 23 Geaster coronatus f. minimus 85 limbatus 90 umbilicatus 84 129

Page  130 130 PUFFBALLS AND ALLIES IN MICHIGAN Geastrum 3, 4, 9, 1o, 49, 8o, 82, 83, 90, 97, 99, 101, 114, 115 asper 84, 97 campestre 84, 96 coronatum 84, 85 fimbriatum 93, 94, 95 f. fimbriatum 84, 92 f. pallidum 84, 94 fornicatum 91 limbatum 84, 90, 94 Morganii 84, 88 pectinatum 84, 86, 88 rufescens 84, 89, 91, 92 saccatum 84, 89, 93, 95, 96 Schmidelii 84, 87, 88 triplex 84, 89, 91, 92, 94, 96 umbilicatum 83, 84 Gellopellis 29 hahashimensis 29 Glischroderma io6 Gymnoglossum 20 Gyrophragmium 123 Holocotylon 13, 18 Hydnangium 21 Hymenogaster 20, 126 niveus 20 Hysterangium 26, 27, 126, 127 clathroides 27 Itajahya 30 Ithyphallus 33 Jaczewskia 26 Jansia 30 Kalchbrennera 30 Lanopila 39 Laternea 30 Le Ratia 18 Leucogaster 13, 15, 126 Leucophleps 13, 15 Longula 123 Lycogalopsis 79 Lycoperdon 3, 7, 9, 11, 39, 45, 46, 47, 56, 58, 59, 62, 64, 66, 70, 73, 74 coloratum 48, 51 Curtisii 47, 50, 56 echinatum 48, 67, 68 gemmatum 58, 59, 6i marginatum 48, 50, 55, 56 muscorum 48, 58 oblongisporum 48, 53, 54 Peckii 48, 59 pedicellatum 48, 6i perlatum 48, 60, 61 polymorphum 48, 51, 55 f. cepaeforme 51, 54 pulcherrimum 42, 48, 66, 67, 68 pusillum 48, 52, 54, 55, 74 pyriforme 48, 49, 56, 61 rimulatum 48, 65 subincarnatum 47, 48, 50 umbrinum 48, 52, 62, 64, 65, 75 var. asterospermum 65 var. atropurpureum 48, 64 var. floccosum 46, 48, 64, 67 var. umbrinum 48, 54, 62, 64, 65 Wrightii 50 Lycoperdopsis 38 Lysurus 30 Maccagnia 15 Macowanites 127 Melanogaster 15, 17, 127 ambiguus 16 Broomeianus i6 Mesophellia 1o, 81 Montagnea 123 Morganella 39 Mutinus 8, 30, 31 caninus 32 var. albus 31, 32, 33 var. caninus 31 Curtisii 31 elegans 31, 32, 33 Mycenastrum 9, 36, 79, 8o corium 80 Myriostoma 82 coliforme 82 Nidula 117 Nidularia 117 Nidulariopsis 121 Pelargonium 21 Phallogaster 26, 28, 126 saccatus 8, 28 Phallus 8, 30, 33 impudicus 11, 33 Ravenelii 33, 34 Phellorinia 107 Pirogaster 98

Page  131 INDEX TO GENERA AND SPECIES 131 Pisolithus 8, 0o5 crassipes 106 tinctorius I05, 106 Podaxis 122, 125 Polytrichum 48, 58 Pompholyx 98 Protubera 29 Pseudocolus 30 Queletia 107 Radiigera 81 Rhizopogon i8, 127 roseolus 19 rubescens 19 Rhodophyllus 20 Rhopalagaster 26 Richoniella 20 Schizostoma 10, 107 Scleroderma 3, 11, 7o, 98, 99, 101, 105 arenicola 99, IOI, 102 aurantium 8, 99, ioo Bovista 102 flavidum 8, 99, IOO, 104 Geaster 8, 99, 103, 104 lycoperdoides 99, o02 vulgare 102 Sclerogaster 21 Secotium 123 agaricoides 123 coprinoides 124 pingui 123 Sedecula 104 Simblum 30 Sphaerobolus 121 iowensis 121 stellatus 121 Sphagnum 48 Staheliomyces 30 Terrostella 82 Torrendia 15 Tremellogaster 25 Trichaster 82 Truncocolumella 18 Tuber 12 Tulostoma 4, 10, I07 brumale 108, III campestre 108, IIO fibrillosum 6, 107, 109, 110 pedunculatum 11 poculatum 109 simulans o18, iII striatum 71, 107, io8, 109 volvulatum 108, 112 Tylostoma 126 mammosum 111 Vaccinium corymbosum loo Xylophallus 30

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PLAJ F. I M e 1 " 71! CLN t T r?ei 7u a B 1 h\ I k- in dG FIG, 2 Alpo:a DodulTL F lk; ' (! " ; t' i '' ''. '. V, I I I! " " I I (-! , i'

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PLA -IE II r-4.K Im _ it.!,. Ar(.-I I P t" I e " , i ", i I.',:,.1 N I () I 2 , , i. -,. 'I Wm 7,-ll ~ S,11FA R

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PLAlIE III (7??Lk Fi cl;2D. i:- fo; I. P;,B>( I'>( j F -1L>

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PLATE IV F7g f lo I r 1.. -.. I 0 S.: 4.. t, )L. I I,-\. 4-, -II. Ix, I " I 5-. IlkP C- I-'.,. Phal"lus Razecneiji Berkelev and Curtis. X 1

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PLATE XI. r., II I " I % I 1~~1, ~ 9 4- -a -v '%I j:, f.,,,. I 4 A.-041, -4 1 - VW We' 'S A0 I c of WII. 4 * k%, I.A N p. D~ic'.1voJ12 C B os c F. sc I i 3.4

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i Ii I i PLATE VI -- - ----— ~ --- ~; -- ------- ~ ~~~- - -~ ----~~ ~- ~ ~ -;~ —~~~, t 1,E*-fiEiBC;a')l -bRY F1Pr-~Eesjaaelif'_ 1 i ',Rcl"rParli;Gs'_L1C;f;%3F. r r r j! 5 i I r B e i m r r r I r I r I r 9 E r I i i B i i j j Ir ~I t a r' S(ILCe j a r 1 O Fra. I. Calvatia cyathifo7mis IBosc.) Morgan. X 1 FIG. 2. Calvatia fragilis (Vitt.) Morgan. X 1

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PLATE VII Z$,,,~e I, Is ":. I Calvatia gigantea (Pers.) Lloyd. X /2

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IILA NI F VII I 1. I..f 'or I! i I I - I #- I,*.-.4 - I. *J& -j" f -l'.w 4rl vv,; " -,?ltt-. "r,. I 1. T - I I -.,,: ,,: -,: " P :,,; r? I; - ( - I \- I! - 1,,

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P1<LA I1F I N. I 1 '4;11. I. j i I I w I I...I q.-.,'Ift I 11,,,.-i '' 1,.N V>,:,' I I I >i

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P LATE X Fi-I LycofpcrdC(l kUbincarniluinl Peck ' AA FIG. 2). Lycoperdon pusillurn Persoon. X I Fe.. Lvcoperdoii oblo)iigil~qoru rn Bcrkele,, and Curtis. /.,. I

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PLATE XLI Fic;. I. Ljcopcirdn-? mar'(7.I 1! II I? Vit t Id IIII i FIG. 2. Lycoperdon mar ginoinut Vittadinji. 1.IiJttitatuir&

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PLATE XII f~~~I\la form" -ItI 1.11.. 1. I I. I I I q 0 *.. " ' ' fli,,, I - 11. '.. 1. I T,,,I I i., r Jtq I~; I I Sc rl -

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.Ilk~I

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I X AI OOSI Ad w it) 1)/ I I,/ 11, /,I ,/,, o -I clhellw AL A..." 11 4(..-, I,:... 4, ' ',,..:, %I.-.. I blll...,.. I " I,11..:,.. 40 I; PI' "..\I \ 'A1 WI \I

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PI AJ YE- x\V I,1 2,, I,~ r,t,,~.,?' P t, - ' aI L - Fc2 Iicocperdon pid(Il('laium?r PeckL'

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I ILAl-I " L XVI JGIL-coperdon2 u mbr ri ai P,,S o o x r. UNu bIn ur m Fi] 2 A' I-:,,!,~, i ~ it1 brZnu'1 Va a! uN O. - )III )I art Uarn ( VIt r. ) I II6 S.

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IT \I I *,V C ".. A I 14 - e, .L -f,,, 1.,.. ,Ff# I I Ir.,,, -,r -, I,, i t- -.;.. i,.. / ,,.. 1.... " " 1,?1; /,! :. i! -. i I i,:,,,,,! !i t' I I;1, )\ k I. I

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PLATE XVIII Lycoperdon umbrinum var. floccosum Lloyd. X 1

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P L.\TF X IX I 1 ~~~~~~~~~ 1, p.~~~~~~~~~ I!,:,, 11 f . - 11!;,, I, "i,. " "I y I (,. I t 1 , I

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PLATE XX FIG. 1. Lycoperdon echinatum Persoon. X 1 I s. S * _ I.. FIG. 2. Lycoperdon echinatum Persoon. X 1

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PLATE XXI, 'J -f, C - _I00-:.WI -; - I FIG. 1. Disciseda candida (Schw.) Lloyd. X 1 ~~~~~~~" 7 l FIG. 2. Disciseda subterranea (Pk.) Coker and Couch. X I

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PLATE XXII I I, IP 7; "on.P;7 touilland (, i LI'tL ' o i 0"

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PLATE XXIII ft.~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~t all ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ f IA4~~~~~~~~~~' I I 2 ~~~~~~\font- P~itoi 4~ r,s(ction ol str l~ ~ Fi 2- Irzsd a?djica N\Iont. 1~Prtoujilli:i d, '-. I Shox:nL 'oit ci 001, c tx

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PLATE XXIV FIG. 1. Bovista pila Berkeley and Curtis. X 1 I FIGo. 2. Bovista plumbea Persoon. X 1 FIG. 3. Bovista minor Morgan. X 1

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PLATE XXV,pC- -.c3L~1".3ad c 3hc 4" "i _zr c, bSi I Y ~ci;r.bP ~. 4P . ib )iF4r~P; r' '~ C'\ ~' -~.h; L ~L c"f~ R:IJe.7 ~,.ii~ ~ b;Sn I, i, ~i 'cY?5: ' tit~' fiF I I c FIG. 1. Mycenastrum corium (Guers.) Desvaux. X Y3 FIGQ.2. Mycenastrum corium (Guers.) Desvaux. > 2/3

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PLATE XXVI FIG. 1. Mycenastrum corium (Guers.) Desvaux. X YA FIG. 2. Geastrum coronatum Persoon. X 1

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PLATEi XXVI I i I -,:, ~ ~I,.~, I ~,? '', j 1, -, I'll V 2 1',I, I -:; X".1 I. kl- ,. A%..1 6 T, V~~~~~~~~~~

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PLATE XXVIII FIG. 1. Geastrum triplex unghuhn. FIG. 1. Geastrum triplex Junghuhn. X 3 FIG. 2. Geastrum triplex Junghuhn. X? FIG. 2. Geastrum triplex Junghuhn. X 3

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PLATE XXIX FIG. 1. Geastrum rufescens P, i ersoon. X 3 FIG. 2. Geastrum rufescens Persoon. X 1 - "V

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i PLATE XXX -ia FIG. 1. Geastrum saccatum Fries. X 1 FIG. 2. Geastrum saccatum Fries. X 3

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PLATE XXXI FIG. 1. Geastrum campestre (Morg.) Kambly and Lee. X 1 I, ~ ~ ~ - fItf l I FIG. 2. Geastrum umbilicatum Fries sensu Morgan. X 1 FIG. 3. Geastrum Morganii Lloyd. X 1

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PLATE XXXII -FIG. 1. Scleroderma aurantium Persoon. X 1 Photograph by E. B. Mains FIG. 2. Geastrum limbatum sensu Coker and Couch. X 1 FIG. 3. Geastrum limbatum. X 1 +

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f W'. I tt oy. 6-Itp A,

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PLATE XXXIV FIG. 1. Scleroderma flavidum Ellis and Everhart. X 1 FIG. 2. Scleroderma flavidum Ellis and Everhart. X 1

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PLATEIF XXXXV F! (. I!t I f, dt 1, i (i Z, 1; it, I. 'I 2 6, ~~~~~~~~~~~~~~~~~~~~~~~~~~~~. f, t

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X AJ X XXVI S~frd1r I(;ea.1' lep, F r1(S. x I

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PLATE XXXVII Pi soI Ith u s!I r CIO 77i7uS PJYS. (. ( )'(-I rl dc ( ou. tICh11-

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PLATE XXXVIII FIG. 1. Tulostoma striatum Cunningham. X 1 t. -_ FIG. 2. Tulostoma fibrillosum White. X 1

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F PLATE XXXIX IG. 1. Tulostoma campestre Morgan. X 2 FIG. 2. Tulostoma simulans Lloyd. X 1 N.O, l 1 1 FIG. 3. Tulostoma simulans Lloyd. X 2 \^

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PLATEi xi. 1 A.;,, r. I, 7 I. I I,,.,, I I . ( I,, C), E, v,, I.,VP a in;.m T, I, I; I i - (I!., Ii% D.( Iw 1\ I

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PLATE XLI Crucibulum levis (D.C.) Kambly. Enlarged. Photograph by E. B. Mains

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PLATE XLII FIG. 1. Sphaerobolus stellatus Persoon. Enlarged Photograph by E. B. Mains FIG. 2. Sphaerobolus stellatus Persoon. X 1

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I %j lll A.:,- 4F,'4,,.,, " Fro.